Scolopendrellidae ( Myriapoda , Symphyla ) from the Afrotropics with descriptions of seven new species

Among the Scolopendrellidae belonging to the collection of Symphyla of the Royal Museum for Central Africa (Tervuren, Belgium), ten species were found and seven new species are described: Symphylella erecta sp. nov., S. fuko sp. nov., S. kalundu sp. nov., S. lubumbashi sp. nov., S. malagassa sp. nov., S. tanganyika sp. nov. and Remysymphyla spinosa sp. nov. SEM micrographs and additional information of the new species and also of Remysymphyla hova Aubry & Masson, 1952, Ribautiella zagnanadina Brölemann, 1926 and R. schoutedeni Hinschberger, 1954 are provided, which contributes to an easier determination of the afrotropical scolopendrellid species. The diagnosis of the genus Remysymphyla is discussed.


Introduction
Little is known of the Symphyla fauna of the Afrotropics, especially regarding the family Scolopendrellidae.Fourteen species of Scolopendrellidae have previously been reported in the Afrotropics.
The present work is a study of the collection of Symphyla of the Royal Museum for Central Africa (Tervuren, Belgium).The vast majority is unidentifi ed material -nearly 300 tubes containing 1700 specimens in ethanol -from the whole Afrotropics and collected from the 1950s until now.Most specimens belong to family Scutigerellidae, a few of them to Scolopendrellidae.
With the present study, the number of now identifi ed species of Scolopendrellidae from the Afrotropics is signifi cantly increased.The study also presents a synopsis of the Afrotropical species belonging to the studied genera.

Material and methods
Twenty eight specimens of scollopendrellids were studied using SEM.Specimens preserved in 70% ethanol were dehydrated in a graded series of acetone up to 100%, critical-point-dried with carbon dioxide in a Balzer CPD 030 and subsequently sputter coated with gold.Morphological characters of the specimens were examined under a JEOL JSM-5400LV microscope at 10 kV.

Etymology
From the Latin erectus = upright, referring to the long and erect setae on the tergites.

Description (holotype)
LENGTH.2.3 mm.HEAD.1.1 times as long as broad with broadest part behind the middle on a level with the points of lateral protuberances (Fig. 1A).Lateral margins nearly straight.Central rod faint, most distinct in posterior half (Fig. 1A).Frontal branches also faint.Median branches absent.Dorsal surface with setae of different lengths.Cuticle with faint and dense pubescence (Fig. 1B).
ANTENNAE.Left antenna 16 segments, right 14 (Fig. 1G).First segment smallest, following segments about 1.2 times as wide as long.Total number of setae visible in dorsal view: 3 on fi rst segment and 4 on second and third ones.Longest setae of proximal segments about 2.3 times as long as those of distal segments.Average length of setae about 0.3 diameter of the basal segments.One whorl of setae on proximal segments, secondary whorl begins on ventral side of 7 th or 8 th segment.Apical segment globular with several short distal setae.All segments with faint pubescence.1A, 1E, 7A).First tergite rudimentary with eight setae arranged in two groups of four.Second tergite complete; triangular posterior processes with straight margins and glabrous tips, no endswelling (Fig. 1A, E).Margin between projections straight.One marginal seta between apical and inner basal setae.Inner basal setae thick, erect and long, as long as anterolateral seta.Eight posteromarginal setae -apical ones included -, two setae in the medial position -central setae.Five lateromarginal setae on each side, -anterolateral setae included -some of them larger and very erect, alternating with the smaller ones.Anterolateral setae very prominent in all tergites except most posteriors, erect, pointing forwards and protruding from both sides.Second tergite with one anterolateral seta on each side, 3rd and following ones with two.Third tergite with six posteromarginal setae and eight lateromarginal on the right side, four visible on the left side -anterolateral setae included -some larger and very erect, alternating with smaller ones.All tergites with faint pubescence.LEGS.First pair of legs not visible in holotype, strongly reduced to small spiny spot with two protruding setae each in paratypes (Fig. 1C, D).Last pair of legs with few and small setae.(Fig. 1H).Tarsus with a few distinct small setae in distal part and two longer and erect setae on dorsal side (Fig 1H,I).Cuticle with very faint pubescence.STYLI.Not visible in holotype, short and subglobular, with short thick hairs in paratypes (Fig. 1F).CERCI.3.2 times as long as wide, slightly curved laterally (Fig. 1I).35-40 setae in dorsal view on each cercus.Setae either short, slightly curved, depressed, clothing all sides, or long straight erect, inserted on outer and ventral sides.Distal half of outer side with 1 or 2 long and erect setae; ventral side with 4-6 erect setae arranged in a longitudinal row (Fig. 1H).Cuticle with faint pubescence.Terminal area conical, 0.1 times of the length of cerci, six-nine transversal striae.Apical seta curved inwards, about as long as terminal area.

Affi nities and differential diagnosis
Symphylella erecta is very similar to S. simplex (Hansen, 1903) and S. multisetosa Scheller, 1971 from South Asia.These species differ from all other Symphylella species in the combination of having 8 setae on the fi rst tergite, setae between the apical and inner basal setae of the triangular processes, some of the lateromarginal setae on the second tergite erect and the presence of at least one row of erected setae in the ventral part of the cerci.However, S. erecta is distinguished by the presence of long inner basal setae on the triangular processes of the second tergite, as long as the largest lateromarginal one, strongly reduced fi rst pair of legs and the presence of frontal branches in the head.The presence of the subglobular styli is also a diagnostic feature of this species.Type material Holotype Sex unkown, CONGO D.R., A.N.P., secteur sud, riv.Rumanura, affl . Fuko, 1200m, coll. De Witte, 14 Jul. 1954 (MRAC 22151).

Etymology
The species name is a noun in apposition taken from the type locality.

Description (holotype)
LENGTH.2.5 mm.HEAD.1.1 times as long as broad with broadest part behind the middle on a level with lateral protuberances (Fig. 2A).Lateral margins nearly straight.Central rod thin, nearly indistinct in the anterior part.Frontal branches present but no median branches.Dorsal surface mostly covered with short setae, the longest are situated in the anterior part and in the lateral margins.Cuticle with faint and dense pubescence (Fig. 2C).ANTENNAE.Left antenna 19 segments, right 18 (Fig. 2G).First segment smaller, 2 nd and 3 rd segments about 1.2 times as wide as long.The following segments are gradually wider, about 1.8 times as wide as long in the 12 th .Setae about 0.2 times the diameter of the segments; shorter in the most distal segments.No second whorl of setae from dorsal view.Apical segment subglobular, slightly wider than long.All segments with faint pubescence.2A, H, 7B).First tergite rudimentary with seven setae arranged in two groups of three and four.Second tergite complete; triangular processes with straight margins and small end-swellings (Fig. 2A, H).Margin between projections straight.One marginal setae between apical and inner basal setae on the left triangular projection, two on the right one; nine posteromarginal setae with equal length, with two central ones; seven lateromarginal setae on each side.Anterolateral setae longer than others.Third tergite with ten lateromarginal setae on the left side, right margin not entirely visible.All tergites with faint pubescence.

TERGITES (Figs
LEGS.First pair of legs strongly reduced to spiny spots with two protruding setae and two longer setae on both sides (Fig. 2B, D).Anterior seta as long as protruding seta of the spot.Posterior seta 4 times longer of anterior one and pointing backwards.Last pair of legs barely visible in the holotype, with three longer seta on dorsal part of the femur -one in the proximal part and two in the distal part -and one on the tibia (Fig. 2F).Cuticle with very faint pubescence.STYLI.Not visible.CERCI.2.5 times as long as wide, sides slightly curved (Fig. 2E).Setae generally short, slightly curved and depressed on all sides; 40 visible from the dorsal view.One erect and longer seta on the ventral side of both cerci (Fig. 2F).Cuticle with very faint pubescence.Terminal area not visible in the holotype.

Affi nities and differential diagnosis
The combination of a relative high number of setae on the fi rst tergite -four setae on the left half -and on the second tergite -nine posteromarginal and seven lateromarginal setae -is shared by S. fuko sp.nov., S. simplex, S. multisetosa and S. neotropica (Hansen, 1903).The presence of only three setae on the right half of the fi rst tergite, longer and erect setae on the second and third tergites and the frontal branches of the central rod on the head are characteristic for S. fuko sp.nov.This species also differs from S. multisetosa and S. neotropica in the fi rst pair of legs, which is strongly reduced to spiny spots.The chaetotaxy and morphology of the fi rst three tergites in S. fuko sp.nov.are also similar to those of S. erecta sp.nov., but S. fuko sp.nov. is easily distinguished by the regular size of the inner basal setae on the second tergite and the shorter anterolateral setae.

Etymology
The species name is a noun in apposition taken from the type locality.

Description (holotype)
LENGTH. 2 mm.HEAD.1.2 times as long as broad, with broadest part behind the middle on a level with the lateral protuberances of the head (Fig. 3A).Lateral margins nearly straight.Central rod distinct, thin, with a node at the middle.Frontal branches present, no median branches.Dorsal surface covered by setae of subequal length.Cuticle with faint and dense pubescence (Fig. 3B).
ANTENNAE.Right antenna broken, left antenna 16 segments (Fig. 3F).First segment elongate, about 0.5 times as wide as long; 2 nd and 3 rd as wide as long; following segments of equal size, about 1.2 times as wide as long.Setae about 0.2 of diameter of segments, shorter in the most distal segments.No second whorl of setae.Apical segment subglobular.All segments with faint pubescence.3A, E, 7C).First tergite rudimentary with six setae arranged in two groups of three.One short seta between fi rst and second tergites in the median part (Fig. 3E).Second tergite with one seta between apical and inner basal setae of the triangular processes.Triangular processes with very small end-swellings (Fig. 3E).Margin between processes slightly curved.Second tergite with a total of seven posteromarginal setae, one central seta in the medial position.Five lateromarginal setae on the right side, short.Anterolateral setae slightly longer than the other lateromarginal setae.Only three lateromarginal setae visible on the left side.Third tergite with fi ve posteromarginal setae, without setae between apical and inner basal setae of the triangular processes, seven lateromarginal setae on the right side, only four visible on the left side.All tergites with very faint, short and sparse pubescence.

TERGITES (Figs
LEGS.First pair of legs reduced to small spiny knobs, each with one distal seta (Fig. 3C, D).Last pair of legs with few and small setae.Tarsus with few long and curved setae in the distal part (Fig. 3G), absent in one paratype (Fig. 3H).Cuticle pubescent.STYLI.Extremely short, barely visible (Fig. 3I).CERCI.3.2 times as long as wide, laterals slightly curved, with 10 dorsal setae long, slightly curved and depressed (Fig. 3G).Ventral side with fi ve protruding and erect setae arranged in a longitudinal row (Fig. 3H).Cuticle pubescent.Terminal area conical, 0.14 times as long as the total length of the cerci, with 4 proximal striae straight and three distal striae curved and concentric.No apical setae, probably lost in all specimens.

Affi nities and differential diagnosis
The species is similar to S. vulgaris (Hansen, 1903) and S. asiatica Scheller, 1971, since they share some similarities in the chaetotaxy of the two fi rst tergites: six setae on the fi rst tergite, seven posteromarginal setae on the second tergite with one central seta in the medial position and one seta between apical and inner basal setae of each triangular process.Further similarities in the chaetotaxy of the tergites are shared by S. kalundu sp.nov.and S. asiatica, such as 5 lateromarginal setae on the second tergite and 8 on the third one, while they are respectively 8 and 10 in S. vulgaris (Hansen, 1903).The absence of setae between the apical and inner basal setae of the triangular processes on the third tergite is characteristic of S. kalundu sp.nov.The chaetotaxy of the cerci is more similar to that of S. vulgaris (Hansen, 1903) than the one observed in S. asiatica, since there is only one row of ventral setae erect, and there are no erect dorsal setae.

Etymology
The species name is a noun in apposition taken from the type locality.

Description (holotype)
LENGTH.1.8 mm.HEAD.1.2 times as long as broad with broadest part in the middle on a level with lateral protuberances (Fig. 4A).Lateral margins nearly straight.Central rod thin, more distinct in the posterior half.Frontal branches not visible in any of the studied specimens.Median branches absent.Dorsal surface with short setae.Cuticle with faint and dense pubescence (Fig. 4B).ANTENNAE.Proportionately short and depressed, left antenna broken, right with 14 segments (Fig. 4F).First segment smaller, following segments pretty broad, about twice as wide as long.Length of setae 0.25-0.3times the diameter of segments, shorter in most distal segments.No second whorl of setae.Apical segment subglobular, wider than long.All segments with faint pubescence.4A, E, 7D).First tergite rudimentary with seven setae arranged in two groups of three and four.Triangular processes of following tergites with straight margins, prominent end-swellings and one seta between apical and inner basal setae (Fig. 4E).One short seta between fi rst and second tergites in the median part (Fig. 4E).Second tergite with seven posteromarginal setae, only one central seta, and four thick and erect lateromarginal setae (Fig. 4E).Anterolateral seta longer and well distinct in all anterior tergites.Margin between triangular projections straight in the second tergite but curved in the following ones.Third tergite with eight posteromarginal setae with two central ones, six-seven lateromarginal setae.Setae of the central part of third tergite clearly shorter than marginal ones.All tergites with a dense and conspicuous pubescence.

TERGITES (Figs
LEGS.First pair of legs strongly reduced to small spiny spot with two protruding setae (Fig. 4C, D).Last pair of legs with protruding setae in the distal part of the joints (Fig. 4H).Femur with one longer distal seta on the dorsal side.Tibia with four protruding setae in the distal part.Tarsus with three thick and protruding distal setae.Cuticle pubescent.STYLI.Small, elongate, covered by hairs resembling a spike (Fig. 4F).CERCI.2.5 times as long as wide, with slightly curved sides (Fig. 4I).25-30 long, straight and depressed dorsal setae.Outer side of distal half with one or two erect setae; ventral side with four or fi ve setae -not erect -arranged in a longitudinal row (Fig. 4H).Cuticle with faint pubescence.Terminal area conical, 0.1 times as long as the total length of the cerci with seven-nine transversal striae (Fig. 4I).Apical seta straight, 1.7 times as long as terminal area.

Affi nities and differential diagnosis
The presence of only four lateromarginal setae on the second tergite is a characteristic feature shared by S. lubumbashi sp.nov., S. foucquei Jupeau, 1954 andS. elongata Scheller, 1952.However, S. lubumbashi sp.nov.differs from the latter two species by the presence of a seta between apical and inner basal setae of the triangular processes on the second and third tergites.

Remarks
There is intraspecifi c variation in the chaetotaxy of the fi rst and second tergites: one paratype presents only six setae on the fi rst tergite, and a small seta between the anterolateral and the following more posterior on the second tergite.

Etymology
The species name is an adjective derived from Madagascar.

Description (holotype)
LENGTH.2.00 mm.HEAD.1.7 times as long as broad with broadest part behind the middle on a level with lateral protuberances (Fig. 5A).Lateral margins of head nearly straight.Central rod only distinct in the posterior half.Frontal branches well developed and median branches indistinct.Dorsal surface with short and thin setae, becoming denser in the posterior part.Cuticle with faint and dense pubescence (Fig. 5B).ANTENNAE.Left antenna with 16 segments, right with 17 (Fig. 5F).First segment small, 0.4 times the length of 2 nd segment.Following segments with regular size, about 1.2 times as wide as long.Inner setae of the second and third segments straight and longer than the ones of the following segments (Fig. 5F).Longest inner seta of the third segment 0.6 times as long as the diameter of segment, while length of the ones in the following segments about 0.3 times the diameter, even shorter in the most distal segments.No second whorl of setae.Apical segment globular, all segments with faint pubescence.5A, E, 7E).First tergite rudimentary with six setae arranged in two groups of three.Following tergites with quite curved lateral margins and well demarcated curved posterior margins between triangular processes.Processes without end-swelling, tips slightly pointing inwards.One seta between fi rst and second tergites in the median part (Fig. 5E).Second tergite with nine posteromarginal, two central and fi ve lateromarginal setae.One seta between apical and inner basal setae on the left triangular process, two on the right one.Anterior margin of second tergite concave.Anterolateral setae longer and well distinct in all anterior tergites.Third tergite with seven-ten lateromarginal and ten posteromarginal setae.

TERGITES (Figs
LEGS.First pair of legs strongly reduced to small spiny spots (Fig. 5C, D).Last pair of legs with small setae on the whole surface.Tarsus with a longitudinal row of fi ve longer setae on the dorsal side (Fig. 5H).Cuticle pubescent.STYLI.Small, elongate and spike-like, covered by hairs, with a club-like distal process (Fig. 5G).
CERCI. 3 times as long as wide, with straight sides (Fig. 5I).25-30 setae in dorsal view.slightly curved and depressed.Cuticle with faint pubescence.Terminal area conical, quite wide, 0.1 times as long as the total length of the cerci, with 8 transversal striae.No apical setae, probably lost in both studied specimens.

Affi nities and differential diagnosis
Symphylella malagassa sp.nov. is similar to S. kalundu sp.nov., S. asiatica Scheller, 1971 andS.brincki Scheller, 1971, since they present six setae on the fi rst tergite and fi ve lateromarginal setae on the second one.S. malagassa sp.nov.differs by the presence of two or three setae between the apical and inner basal setae of the triangular processes instead of one, the higher number of posteromarginal setae on the third tergite and the characteristic curvature of the posterior margin of the tergites.Another remarkable diagnostic feature of this species is the club-like distal end of the styli.Symphylella tanganyika sp.nov.urn:lsid:zoobank.org:act:E92B089D-B6F7-44C2-8CC1-52CDFA5BE742Figs 6, 7F Type material Holotype Sex unknown, TANZANIA, Mt.Uluguru, Kinola, 1500-1750 m, coll.Berger, Leleup and Debecker, 6-13 Jun. 1971 (MRAC 22159).

Etymology
The species name is a noun in apposition taken from the old name of the type locality.

Description (holotype)
LENGTH.1.7 mm.HEAD.1.2 times as long as broad with broadest part behind the middle on a level with lateral protuberances (Fig. 6A).Lateral margins rounded.Central rod very thin, well distinct in the posterior part.Frontal branches present, faint, and median branches absent.Dorsal surface with rather short setae.Cuticle with faint and dense pubescence (Fig. 6B).ANTENNAE.Left antenna with 16 segments, right with 17 (Fig. 6F).First segment much smaller than following ones, about 0.3 times the length of 2 nd segment.Following segments with regular size.Inner setae of 2 nd and 3 rd segments longer than the ones of the following segments.Longest inner seta of the third segment 0.4 times the diameter of segment, length of the ones in the following segments about 0.2-0.3times of diameter, gradually shorter towards the apex.Secondary whorl of setae begins on the ventral side of the 7 th segment.Apical segment globular, with several small terminal setae.All segments with faint pubescence.6A, E, 7F).Small in comparison with the general width of the head and the body; width 0.6-0.8times of the diameter of the trunk at the same level.First tergite rudimentary with eight setae arranged in two groups of four.One seta between fi rst and second tergites in the median part (Fig. 6E).Lateral and posterior margins of second tergite straight.Nine posteromarginal setae with two central setae.One seta between apical and inner basal setae on the left triangular process, two on the right one.Six lateromarginal setae.Anterolateral setae comparatively not very long in the anterior tergites.Third tergite with eight posteromarginal setae and nine lateromarginal.Setae of the central part of third tergite clearly shorter than marginal ones.All tergites with faint pubescence.

TERGITES (Figs
LEGS.First pair of legs strongly reduced, in a small spiny spot with two protruding setae (Fig. 6C,  D).Last pair of legs with thick and erect setae, sparse in the dorsal parts of the joints: femur with one erect seta in the distal part, tibia with two and tarsus with three protruding setae in a longitudinal row (Fig. 6H).Cuticle pubescent.STYLI.Very small, subglobular, covered by a moderate number of spiny hairs (Fig. 6G).CERCI.2.5-2.7 times as long as wide, slightly curved laterally (Fig. 6I) and dorsally straight (Fig. 6H).Many short, slightly curved and depressed setae on all sides.Outer side with one protruding setae in the distal part (Fig. 6I), ventral with fi ve setae arranged in a longitudinal row (Fig. 6H).Terminal area of the right cercus not entirely visible; terminal area of the left cercus conical, quite wide, 0.1 times as long as the total length of the cerci, with fi ve transversal proximal striae and four curved, concentric distal striae (Fig. 6I).Apical seta curved, about as long as terminal area.

Affi nities and differential diagnosis
S. tanganyika sp.nov. is similar to S. erecta sp.nov.; they share the presence of eight setae on the fi rst tergite and the subglobular styli.It is distinguished from S. erecta sp.nov. the chaetotaxy of the second and third tergites, the length of the antennae, which are longer in S. tanganyika sp.nov., the smaller general size of the tergites and the presence of several protruding setae on the dorsal side of the last pair of legs.Edwards 1959).However, the observation of type material of Remysymphyla hova Aubry & Masson, 1952b (MNHN K002) using SEM has shown that it really presents 21 tergites.It means that the tergites of the tergal areas 6, 9, 12, 13, 14 and 15 (fi de Domínguez Camacho 2009: 108, fi g. 4) are subdivided into two subtergites each (Figs 8A, E, 10A), as occurs in many scolopendrellid genera.

Genus
Tergites of the tergal areas 3-14 (fi de Domínguez Camacho 2009: 108, fi g. 4) partially sclerotized; only both exterior parts of these tergites are completely sclerotized (Fig. 8A, E).This diagnostic character has to be verifi ed carefully, since it has been observed only in two species.In Remysymphyla spinosa sp.nov. the second tergite is partially sclerotized (Fig. 9A, E).
Last tergite subelliptical, with posterior margin convex ( Figs 8E, F, 10B).In addition, the size of the last tergite is smaller than in Symphylella and its shape resembles Ribautiella.
Terminal area of cerci smaller than in Symphylella, with diagonal striae rather than transversal, pointing back-and inwards (Fig. 8F, 10B).Basis of terminal area also diagonal back-and inwards rather than transversal -as in Symphylella.Apical seta also pointing inwards (Fig. 10B, C).The shape of the terminal area of cerci also resembles in some aspects Ribautiella.This character has to be observed carefully since it has been inferred from diagrams in the literature (Aubry & Masson 1952a, 1952b;Scheller 1971) and directly observed only in two species.

Remarks
In addition to the low number of described species of this genus, the material studied and cited in the literature is also very scant.Therefore, the gross incongruence detected between the studied material in this work and the descriptions in the literature -specifi cally regarding Remysymphyla hova -leads the authors to think that the presence of only 15 tergites is a dubious autapomorphy.It was cited for the fi rst time in the defi nition for the genus (Aubry & Masson 1952a) but it was not accompanied by diagrams or detailed descriptions of this feature, neither in further works on other species.For this reason, the authors draw attention to this feature for future studies on this genus, since the number of tergites has a major role in the systematics and phylogeny of the scolopendrellid symphylans (Edwards 1959;Domínguez Camacho 2009).Type locality CONGO D.R., Kivu, Itombwe, terr.D'Uvira, tête de source de la Nyalengwe (03º30' S, 029º00' E), in a mountain forest with bamboo, 2500 m.

Etymology
The species name is an adjective emphasizing the long and erect setae of the tergites, which give this species a spiny aspect.

Description (holotype)
LENGTH.2.1 mm.HEAD.1.3 times as long as broad with broadest part behind the middle on a level with lateral protuberances (Fig. 9A).Lateral margins nearly straight.Central rod well-developed and not broken.Frontal branches also well-developed.Median branches not very distinct in the holotype, but clearly distinct in the paratypes (Fig. 9B).Dorsal surface with very long and spiny setae of equal length.Cuticle with dense granulation (Fig. 9G).ANTENNAE.Broken in holotype.Rather short, in paratype left antenna with 27 segments, right with 29, (Fig. 9C).All segments of equal length and pretty broad, 1.8-1.9times as wide as long.Antennae densely setose, with a spiny appearance; setae erect and long.Longest setae in the inner part of the median segments, with length of 0.5-0.7 times the diameter of segments.Secondary whorl of setae appears in the 6 th -8 th segments.Size and number of setae decreases gradually in the seven distal segments, and are totally absent in the last three ones.Apical segment fl attened, more than 3 times as wide as long.It presents a transversal depression, bestowing a coffee-bean shape.All segments with pretty long and conspicuous pubescence.
TERGITES.Not all well-visible in the studied specimens, at least 17 can be counted in the holotype, of which 13 present triangular processes (Figs 9A, 10A).All tergites -except the fi rst and the two latestsclerotized only in the lateral parts.All tergal setae very long; shortest setae at least as long as triangular processes.Triangular processes longer than broad in the anterior tergites, with digitiform ends and no apical setae; processes of the posterior tergites thinner and reduced to small digitiform projections (Fig. 10C).First tergite with six setae arranged in two groups of three.Second tergite comparatively very large (Fig. 9A, E), with fi ve posteromarginal, one central and six lateromarginal setae.Anterolateral setae not longer than the rest.All tergites with pubescence.Last tergite subelliptical, with posterior margin convex (Fig. 10B).
LEGS.First pair of legs reduced to two oviform knobs densely covered by numerous spines (Fig. 9F, H).Two longer setae on the laterals, about twice as long as the whole knob.Last pair of legs with thick and long setae at the distal part of the joints (Fig. 10B, C).Femur and tibia with three setae; tarsus with two protruding dorsal setae.Cuticle pubescent.STYLI.Small, elongate and spike-like, covered by hairs (Fig. 9D).CERCI.2.8 times as long as wide, with straight margins (Fig. 10B, C).About 25 visible setae from the dorsal view, slightly curved and depressed.Cuticle with faint pubescence.Limit between cerci sensu stricto and terminal areas not transversal, but diagonal, as a continuation of the outer margin towards the inner margin (Fig. 10B).Terminal area small, 0.1 times as long as the total length of the cerci, conical, longer in the outer margin than in the inner with six diagonal striae, parallel to the basis of the terminal area.Apical seta about as long as terminal areas, curved inwards (Fig. 10B, C).

Affi nities and differential diagnosis
The morphology of the fi rst pair of legs, which does not present evidence of several joints, makes Remysymphyla spinosa sp.nov.more similar to R. hebetocornuta Scheller, 1971 than to the remaining African species.The spiny aspect of the body is characteristic for this species.

Remarks
The genus attribution of this species has been problematic, mainly due to the above-explained confusion regarding the defi nition of Remysymphyla.The study of type specimens of Remysymphyla hova Aubry & Masson, 1952 allowed the recognition of the fi rst pair of legs of R. spinosa sp.nov.as typical for this genus (Figs 8B-D; 9F, H).However, observations using SEM seems to be suboptimal for detection of one of its major diagnostic characters: the number of joints on the fi rst pair of legs.Although no joints were discernible in this species, the shape of the oviforms knobs of R. spinosa sp.nov.resembles the two-jointed legs of R. hebetocornuta Scheller, 1971.Genus Ribautiella Brölemann, 1926.

Diagnosis
23 tergites, which means that the tergal areas 2, 3, 6, 9, 12, 13, 14 and 15 (fi de Domínguez Camacho, 2009, fi g. 4, page 108) are subdivided into two subtergites; fi rst tergite small, tergites 3, 5-7, 9-11, 13-15, 17, 18 and 21 with paired, triangular posterior processes.In his Keys to the Genera of Symphyla, Edwards (1959) mentions the presence of 24 tergites as a diagnostic feature for this genus.This higher number is due to a supposed subdivision of the fi rst tergite.However, this characteristic is questionable since descriptions and illustrations are not satisfactory and this subdivision appears in different grades among the species.The present study using SEM suggests that it is probably an artefact of the mounts for light microscopy (Fig. 11A, D).
First pair of legs reduced either to hairy or spiny knobs or spots.This diagnostic character is also present in Symphylella.Accordingly, the combination of the characters 1 and 2 is the strongest diagnostic character set for Ribautiella.
Terminal area of cerci small, with basis diagonal, pointing back-and inwards rather than transverse (Fig. 11B, E).Striae -if present -also diagonal rather than transverse, pointing back-and inwards (Fig. 11F).

Ribautiella zagnanadina
Abbreviations A.N.P. = Albert National Park (Democratic Republic of the Congo) Congo D.R. = Democratic Republic of the Congo MNHN = Muséum national d'Histoire naturelle, Paris MRAC = Musée Royal de l'Afrique centrale (Royal Museum for Central Africa) SEM = scanning electron microscopy

Fig. 2 .
Fig. 2. Symphylella fuko sp.nov., holotype.A. Head and fi rst 4 tergites (T1, T2, T3, T4), dorsal view.Cr, central rod; fb, frontal branches; lp, lateral protuberance of head.B. Head and anterior part of trunk, lateral view.C. Detail of the surface of head and lateral protuberance of head (lp).D. Detail of fi rst leg.E. Posterior part of trunk and cerci (C), dorsal view.F. Posterior part of trunk, last right legs and cerci (C), lateral view.Arrows, longer and erect setae of femur (f) of last leg; es, erect setae of ventral side of cerci.G. Left antenna.H. Detail of fi rst (T1) and second (T2) tergites.Arrows, small end-swellings of triangular processes; al, anterolateral setae; as, apical setae; cs, central setae of the posteromarginal setae; ibs, inner basal setae of the posteromarginal setae.

Fig. 3 .
Fig. 3. Symphylella kalundu sp.nov., holotype (dorsal views) and paratype (ventral and lateral views).A. Head and fi rst 4 tergites (T1, T2, T3, T4), dorsal view.Arrow: node in the middle of the central rod (cr); fb, frontal branches, lp, lateral protuberance of the head.B. Detail of the surface of the head and lateral protuberance (lp).C. Detail of the fi rst leg.D. Head and anterior part of the trunk, ventral view.E. Detail of the fi rst (T1) and second (T2) tergites.Arrows, small end-swelling of triangular processes; as, apical setae; cs, central seta of the posteromarginal setae; ibs, inner basal setae of the posteromarginal setae; s, seta between fi rst and second tergites.F. Right antenna.G.Last tergite, last pair of legs and cerci (C), dorsal view.Arrows, longer and erect setae of tarsus (t) of last pair of legs; ta, terminal area of cerci.H. Posterior part of the trunk, last right leg and right cercus (C), lateral view.Es, erect setae of ventral side of cerci; t, tarsus of last leg, without setae.I. Stylus of the leg 10 (arrow and circle).

Fig. 4 .
Fig. 4. Symphylella lubumbashi sp.nov., holotype (dorsal views) and paratype (ventral and lateral views).A. Head and fi rst 4 tergites (T1, T2, T3, T4), dorsal view.Cr, central rod; lp, lateral protuberances of the head.B. Detail of the surface of the head.C. Detail of the fi rst leg.D. Head and fi rst 2 pairs of legs, ventro-lateral view.E. Detail of fi rst (T1) and second (T2) tergites.Arrows, small end-swelling of triangular processes; al, anterolateral setae; as, apical setae; cs, central seta of the posteromarginal setae; ibs, inner basal setae of the posteromarginal setae; s, seta between fi rst and second tergites.F. Right antenna.G. Stylus of the leg 11 (arrow).H. Posterior part of the trunk, last pair of legs and cerci (C), lateral view.White arrows, protruding setae of the femur (f), tibia (t) and tarsus (ta) of last left leg; grey arrows, protruding setae of tibia (t) and tarsus (ta) of the right left; vs, ventral setae of right cercus, in a row.I.Last tergite and cerci (C), dorsal view.As, apical setae; es, erect setae of outer side of cerci, ta, terminal area.

Fig. 5 .
Fig. 5. Symphylella malagassa sp.nov., holotype (dorsal views) and paratype (ventral and lateral views).A. Head and fi rst 4 tergites (T1, T2, T3, T4), dorsal view.Cr, central rod; fb, frontal branches; lp, lateral protuberances of the head.B. Detail of the surface of the head.C. Detail of the fi rst leg.D. Head and fi rst 2 pairs of legs, ventro-lateral view.E. Detail of fi rst (T1) and second (T2) tergites.Al, anterolateral setae; as, apical setae; cs, central setae of the posterolateral setae; ibs, inner basal setae of the posteromarginal setae; s, seta between fi rst and second tergites.F. Right antenna.Arrows, longer inner setae of second and third segments.G. Stylus of the leg 11. H. Posterior part of the trunk, last pairs of legs and cerci (C), lateral view.Arrows, longer setae of the tarsus (t) of last left leg.I.Last tergite and cerci (C), dorsal view.Ta, terminal area.

Fig. 6 .
Fig. 6.Symphylella tanganyika sp.nov., holotype (dorsal views) and paratype (ventral and lateral views).A. Head and fi rst 4 tergites (T1, T2, T3, T4), dorsal view.Cr, central rod; fb, frontal branches; lp, lateral protuberances of the head.B. Detail of the surface of the head and lateral protuberance (lp).C. Detail of the fi rst leg.D. Head and fi rst 2 pairs of legs, ventral view.E. Detail of fi rst (T1) and second (T2) tergites.Arrows, small end-swelling of triangular processes; al, anterolateral setae; as, apical setae; cs, central seta of the posteromarginal setae; ibs, inner basal setae of the posteromarginal setae; s, seta between fi rst and second tergites.F. Left antenna.Arrows, longer inner setae of the second and third segments.G. Stylus of the leg 11 (arrow).H. Posterior part of the trunk, last pairs of legs and cerci (C), lateral view.Arrows, longer setae of the femur (f), tibia (t) and tarsus (ta) of last legs; es, erect setae of ventral side of right cercus.I.Last tergite and cerci (C), dorsal view.As, apical setae; es, erect setae of outer side of cerci; ta, terminal area.

Fig. 9 .
Fig. 9. Remysymphyla spinosa sp.nov. A. Head and fi rst 3 tergites (T1, T2, T3) of holotype, dorsal view.Arrows, digitiform end of triangular processes; al, anterolateral setae; cr, central rod; fb, frontal branches; lp, lateral protuberance of the head; mb, median branches.B. Head and fi rst tergite (T1) of paratype, dorsal view.Cr, central rod; lp, lateral protuberance of the head; mb, median branches.C. Antennae of paratype.Arrows, transversal depression of apical segments.D. Stylus of the leg 11 (arrow) of paratype.E. Detail of fi rst (T1) and second (T2) tergites of holotype.Arrows, digitiform end of triangular processes; al, anterolateral seta; cs, central seta.F. Head and fi rst legs of paratype, lateral view.G. Detail of the surface of the head of paratype.H. Detail of the fi rst leg of paratype.