European Journal of Taxonomy 150: 1-25

http://dx.doi.org/10.5852/ejt.2015.150

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ISSN 2118-9773

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2015 • Enghoff H. & Frederiksen S.B.

Research article

urn:lsid:zoobank.org:pub:D546A676-35F8-4025-8660-F123A0BA67F7

A mountain of millipedes II: The genus Aquattuor

Frederiksen, 2013 - five new species from the Udzungwa

Mountains and one from Mt. Kilimanjaro, Tanzania

(Diplopoda, Spirostreptida, Odontopygidae)

Henrik ENGHOFF 1 * & Sara B. FREDERIKSEN 12

Zoological Museum, Natural History Museum of Denmark, University of Copenhagen,

Universitetsparken 15, DK-2100, Copenhagen 0, Denmark

2 Department of Ecology, Animal Ecology, University of Marburg, Karl-von-Frisch-Strafie 8,

D-35032 Marburg, Germany

* Corresponding author: [email protected]

2 Email: [email protected]

* urn:lsid:zoobank.org:author:FB09A817-000D-43C3-BCC4-2BClE5373635

2 urn: lsid: zoobank, org: author: 6DE62DD5-0055-424A-A13 6-0EF290AF5 AFP

Abstract. The genus Aquattuor Frederiksen, 2013 is revised. A. denticulatus Frederiksen, 2013 (type

species) from the East Usambara Mts, Tanzania, is redescribed, and six new species are described:

A. daudiahempae sp. nov. from Mt. Kilimanjaro, Tanzania, and five species from the Udzungwa Mts,

Tanzania: A. longipala Enghoff sp. nov., A. major Enghoff sp. nov., A. stereosathe Enghoff sp. nov.,

A. submajor Enghoff sp. nov., and A. udzungwensis Enghoff sp. nov.

Keywords. Eastern Arc, Kilimanjaro, taxonomy, new species.

Enghoff H. & Frederiksen S.B. 2015. A mountain of millipedes II: The genus Aquattuor Frederiksen, 2013 - five

new species from the Udzungwa Mountains and one from Mt. Kilimanjaro, Tanzania (Diplopoda, Spirostreptida,

Odontopygidae). European Journal of Taxonomy 150: 1-25. http://dx.doi.org/10.5852/eit.2015.15Q

Introduction

This is the second in a series of articles about the millipedes, especially the family Odontopygidae, of

the Udzungwa Mountains, Tanzania. For general information on the Odontopygidae and the Udzungwa

Mountains, see the first paper in the series (Enghoff 2014). The present article, although not numbered

as such, also forms a sequel to our previous papers on the odontopygids of East Africa (Frederiksen

2013a, 2013b; Frederiksen & Enghoff 2012, 2015).

The genus Aquattuor was described by Frederiksen (2013b) for a relatively small odontopygid from

the East Usambara Mountains, Tanzania. The genus, which has remained monotypic until now, is

characterized by a slender, whiplike gonopodal solenomere and a simple telomere, combined with a

unique structure of the metazonital limbus which consists of large, rectangular, easily detached flaps.

European Journal of Taxonomy 150 : 1-25 ( 2015 )

Collecting activities in the Udzungwa Mountains by the NGO Frontier Tanzania and by staff and

students from the University of Copenhagen (cf Enghoff 2014) have resulted in a considerable amount

of material containing several new species of Aquattuor. At the same time, collections made in the

framework of The KiLi Project ( https ://www. kil im ani aro.biozentrum. un i-wuerzburg. de/ F sponsored by

the German Research Foundation ( cf. Frederilcsen & Enghoff 2015), have procured several samples of

still another new species.

In the present article we describe six new species of Aquattuor , re-describe the type species,

A. denticulatus Frederiksen, 2013, and provide an updated definition of the genus as well as a key to its

species.

Material and methods

Forreasons explained by Enghoff (2014), only adult males were considered. Atotal of 85 adult Aquattuor

males were examined. All specimens are stored in 70% ethanol and are kept in the Natural History

Museum of Denmark, University of Copenhagen (ZMUC). Specimens were examined in 75% ethanol

under a stereo microscope. Numbers of podous rings were counted and midbody vertical diameter

measured on complete specimens. Body parts for scanning electron microscopy (SEM) were transferred

to 96% ethanol, then to acetone, air-dried, mounted on aluminium stubs or on triangles of flexible

aluminium tape and in turn mounted on stubs, coated with platinum/palladium and studied in a JEOL

JSM-6335F scanning electron microscope.

A general description of the genus is provided. The species diagnoses include, inter alia , information

on the number of body rings and body diameter. Although it cannot be excluded that specimens falling

outside the indicated ranges may be found, this information nevertheless helps to characterize each

species. Gonopod terminology is as far as possible as in Enghoff (2014). Because of the great uniformity

of Aquattuor species, and because size information and gonopod details are included in the diagnosis,

the traditional “Description” paragraph has been replaced by a short note on colouration. Adding more

would be redundant.

For the species from the Udzungwa Mts, species of Aquattuor and Chaleponcus (cf Enghoff 2014)

occurring in the same samples and localities are listed under “coexisting species”.

Abbreviations for morphological terms used in descriptions and on illustrations of gonopods

ba =

basomere

bs =

basomeral spine

btl =

basal telomeral lamella

li =

lateral incision of coxa

mbl =

meso-basal lobe of palette

mi =

mesal incision of coxa

mp =

metaplica

mpl =

mesal-posterior lamella of telomere

mpr =

metaplical ridge

pa =

palette

pn =

posttorsal narrowing

PP =

proplica

prl =

proplical lobe

slm =

solenomere

tm =

telomere

triangular telomeral tooth

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Abbreviations used in the text, other than abbreviations for morphological terms

asl = above sea level

FR = Forest Reserve

ZMUC = Zoological Museum, Natural History Museum of Denmark

Results

Class Diplopoda Blainville-Gervais, 1844

Order Spirostreptida Brandt, 1833

Family Odontopygidae Attems, 1909

Subfamily Archepyginae Manfredi, 1939

Tribe Prionopetalini Hoffman, 1991

Genus Aquattuor Frederiksen, 2013

This genus was established by Frederiksen (2013b) for the type and until now only species, A. denticulatus

(East Usambara Mts, Tanzania). The name is regarded as a masculine noun.

Diagnosis

A genus of Odontopygidae Prionopetalini in which the gonopodal proplica is apically expanded into

a distal ‘palette’, the metaplica has an oblique flange, the solenomere is simple, thin and whiplike, the

telomere terminates in a long, curved gutter-like to tubular part, and the limbus of the body rings consists

of large, rectangular, easily detached flaps.

Etymology

The genus owes its name to the unique limbus flaps which resemble tiny sheets of paper in the standard

A4 format.

Description

This description only applies to males and, as far as non-gonopodal characters are concerned, only

includes a selection.

Non-gonopodal characters

Body length. 19-35 mm Midbody vertical diameter 1.4-2.1 mm. 44-54 podous rings, no apodous

rings in front of telson. See Fig. 2.

Colour. Colours of specimens from Udzungwa and East Usambara Mountains all partly or completely

faded, leaving only newly collected specimens from Mt. Kilimanjaro with fresh colour (Fig. 1). Fight

mid-dorsal longitudinal stripe or traces thereof sometimes present, even on faded specimens.

Fimbus. Consisting of relatively large (c. 0.04 x 0.06 mm) rectangular flaps which are easily detached

(Fig. 3D-E).

Telson (Fig. 3B-C). Preanal ring with wrinkled/coriaceous sculpture dorsally. Anal valves with

wrinkled/coriaceous sculpture, with a pronounced dorsal denticle and a smaller ventral one, denticles

well set off from rest of valve rather than just being “sharp corners”. Each valve with three setae. Free

margin (“lip”) of anal valves raised and provided with three small tubercles on which the setae are borne.

European Journal of Taxonomy 150 : 1-25 ( 2015 )

Gonopods (see Fig. 4)

Coxa. In anterior or posterior view 3 V2-AV2 x as long as broad. Margins of proplica (pp) parallel or

slightly diverging in basal c. Vs, apical c. Vs set off by mesal (mi) and lateral (//) incisions; apical part

of proplica appearing like a rounded “palette” (pa). Proplical lobe (prl) protruding into mesal incision.

Metaplica (mp) with irregular surface; at level of mesal and lateral incisions a stout ridge (mpr) running

obliquely across metaplica.

Telopodite. Basomere (ba) with small spine (bs) just distal oftorsotope (Fig. 11 A, C), arculus 90°. Post-

torsal narrowing (pn) pronounced (e.g., Figs 5A, 6A). Telopodite beyond pn divided into solenomere

(slm) and telomere (tm).

Solenomere. Long, thin, whiplike, often resting within telomeral gutter but on preserved specimens

frequently free (probably an artefact), apical part with oblique-longitudinal flutings.

Telomere. At base expanded into a complicated basal telomeral lamella ( btl ) obscuring origin of soleno¬

mere (F ig. 5D). Main part of telomere consisting of a long, narrow lamella folded into a more or less narrow

gutter, sometimes even tube-like in distalmost part; gutter/tube describing a 90°-360° curve, usually in

almost one plane, but sometimes (A. stereosathe Enghoff sp. nov.) clearly in three dimensions (note: this

applies to preserved specimens). Posterior margin of gutter expanded into mesal-posterior lamella (mpl)

(e.g., Fig. 5D). Tip of telomere variable: margins smooth or denticulate-laciniate, internal surface of

gutter more or less microspiculate (e.g.. Fig. 6C-D).

Distribution

Species of Aquattuor are so far known only from Tanzania. They have been found in several mountain

blocks belonging to the Eastern Arc Mts (Burgess et al. 2007): Udzungwa (5 species), Nguru

(1 species), Rubeho (1 species) and East Usambara (1 species). One further species has been found on

Mt. Kilimanjaro. For an overview, see Figs 12-13.

Fig. 1. Aquattuor claudiahempae sp. nov., two paratype S<3 after 18 months in alcohol. Scale bar =

1 mm (B). Photographs by S. Reboleira.

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Intercalary cuticular microscutes

In A. submajor Enghoff sp. nov. and A. udzungwensis Enghoff sp. nov., intercalary cuticular microscutes,

similar to those first reported by Akkari & Enghoff (2011) in polydesmidan millipedes and subsequently

found, inter alia , in the odontopygid genus Chaleponcus Attems, 1914 (Enghoff 2014) and the

callipodidan genus Lusitanipus Mauries, 1978 (Reboleira & Enghoff 2015), were observed.

Notes on identification

The species of Aquattuor are all quite similar, even in gonopod structure. In this respect, they form a

contrast to the previously treated (much larger) species-swarm centred in the Udzungwa Mts., i.e., the

Chaleponcus dabagaensis-growy (Enghoff 2014), in which gonopods are highly diverse. Differences

between Aquattuor species concern body size (Fig. 2), details of the outline of the gonopod coxa, the

curvature of the telomere, the development of the posterior margin of the telomere, and the armature of

the telomere tip.

Similar genera

By far the most diagnostic character of Aquattuor is the large (sub)rectangular-lobed limbus. Sub-

rectangular limbus lobes do occur in certain other odontopygid genera, e.g., Allantogonus Attems, 1912

(Kraus 1960: fig. 5) and Syndesmogenus Attems, 1909 (Kraus 1966: figs 97-99), but in these genera, the

lobes are much smaller and are not prone to detachment. The gonopods of Aquattuor , with their long,

whip-like solenomere and slender telomere, superficially resemble those of Allantogonus (Kraus 1960:

2.2

2 .1

"re

1,9

in lj W

-C

■C

1,7

1,6

1,5

□

Olun.gipala

□ stereosathe

A udrunR.wHnsi5

A major

O sub major

• da Lidia hem pae

X denticulate

1/4

46 48

Number of podous rings

Fig. 2. Body size (numbers of podous rings and midbody vertical diameter) in $$ of species of

Aquattuor Frederiksen, 2013. In cases of (almost) coinciding values, symbols have been slightly

displaced horizontally.

European Journal of Taxonomy 150 : 1-25 ( 2015 )

Fig. 3. Aquattuor spp. A. A. stereosathe Enghoff sp. nov., paratype, front end. B-C. A. udzungwensis

Enghoff sp. nov., paratype, telson, lateral and posterior view. D-E. A. udzungwensis Enghoff sp. nov.,

limbus. D. Specimen from West Kilombero Scarp FR. E. Paratype. Scale bars: A-D = 0.1 mm; E = 0.01

mm.

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

figs 7, 11, 17) and Lamelloramus Frederiksen, 2013 (Frederiksen 2013: figs 5, 9). However, the coxal

apex in Allantogonus is folded basad over the anterior side; the metaplical flange of Lamelloramus is

situated completely differently on the median side, and the telomere is either strongly curling up or

looping in these genera. The limbus is completely different from that of Aquattuor in both.

Included species (alphabetically)

Aquattuor claudiahempae Enghoff & Frederiksen sp. nov.

A. denticulatus Frederiksen, 2013 (type species)

A. longipala Enghoff sp. nov.

A. major Enghoff sp. nov.

A. stereosathe Enghoff sp. nov.

A. submajor Enghoff sp. nov.

A. udzungwensis Enghoff sp. nov.

Species from the Udzungwa Mountains

Aquattuor submajor Enghoff sp. nov.

urn:lsid:zoobank.org:act:690C121A-D087-400D-95BC-42BCA7B224BE

Figs 4-5

Diagnosis

Diameter 1.55-1.87 mm, 45—48 podous rings. Gonopods: lateral incision of coxa (//) absent to poorly

developed; apical palette (pa) rounded, not longer than broad; telomere describing a c. 270° curve in

almost one plane; mesal-posterior lamella of telomere (mpl) low; anterior margin of telomere without a

triangular tooth; telomere tip with finely dentate margins, internal surface microspiculate.

Etymology

The name is an artificial Latin adjective meaning “below major ” and refers to 1) the smaller size of this

species compared to A. major Enghoff sp. nov. and 2) the fact that this species occurs at lower altitudes

than A. major Enghoff sp. nov.

Material studied (total: 17 SS)

Holotype

TANZANIA: S, Morogoro Region, Udzungwa Mts, Mwanihana FR, above Sanje, 1000 m asl, 18 Aug.

1982, in log, M. Stoltze & N. Scharff leg. (ZMUC).

Paratypes

TANZANIA: 7 SS, same data as holotype, but in litter, log and pitfall trap, M. Stoltze & N. Scharff leg.

(ZMUC); 1 S, same data but 1100 m asl, pitfall trap (ZMUC); 8 $$, same data but 1250 m asl, 1 Aug.

1982, under log, M. Stoltze & N. Scharff leg. (ZMUC).

Type locality

TANZANIA: Morogoro Region, Udzungwa Mts, Mwanihana FR, above Sanje, 1000 m asl.

Colouration

After 33 years in alcohol faded, only some darker markings on head and collum left. No traces of a mid¬

dorsal stripe.

European Journal of Taxonomy 150 : 1-25 ( 2015 )

Fig. 4. Aquattuor submajor Enghoff sp. nov., paratype. A. Gonopods in situ , right gonopod coloured.

B. Left gonopod coxa and basal part of telopodite. C. Left gonopod telopodite and part of coxa, dorsal

(basal) view. Articifical colours: orange = coxa; blue = basomere; yellow = solenomere; pink = telomere;

red = gonopod sternum; light green = ventral part of eighth pleurotergite (right side); dark green =

sternum of reduced ninth leg-pair, mp = metaplica; pa = palette; pp = proplica. Scale bars = 0.1 mm.

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Fig. 5. Aquattuor submajor Enghoff sp. nov., paratype, left gonopod. A. Anterior view. B. Close-up of

mesal incision. C. Tip of telomere. D. Posterior view, btl = basal telomeral lamella; li = lateral incision;

mi = mesal incision; mpl = mesal-posterior lamella of telomere; pa = palette; pn = posttorsal narrowing.

Scale bars: A, D = 0.1 mm; B-C = 0.01 mm

European Journal of Taxonomy 150: 1-25 (2015)

Distribution and habitat

Known only from Mwanihana FR. Altitudinal range: 1000-1250 m asl.

Coexisting species

Aquattuor major Enghoff sp. nov. and Chaleponcus mwanihanensis Enghoff, 2014 occur in the same

area, but only at higher altitudes.

Aquattuor major Enghoff sp. nov.

urn:lsid:zoobank.org:act:7DE65287-B22D-4470-A2AF-70556B77DEE9

Fig. 6

Diagnosis

Diameter 1.97-2.11 mm, 53-54 podous rings. Gonopods: lateral incision of coxa ( li ) triangular, basally

delimited by triangular tooth; apical palette (pa) obliquely rounded, much longer than broad; telomere

describing an almost 360° curve in almost one plane; mesal-posterior lamella of telomere (mpl) low;

anterior margin of telomere with a triangular tooth (tt) ; telomere tip with densely microspiculate margins

and internal surface.

Etymology

The name is a Latin adjective meaning “larger” and refers of the larger size of this species, compared to

its congeners.

Material studied (total: 4 ff )

Holotype

TANZANIA: Morogoro Region, Udzungwa Mts, Mwanihana FR, 1800-1850 m asl, 25-29 Nov.

1984, pitfall traps, montane rain forest, N. Scharff leg. (ZMUC).

Paratypes

TANZANIA: 3 SS, Morogoro Region, Udzungwa Mts, Mwanihana FR, above Sanje, 1650 m asl,

18 Aug. 1982, pitfall trap, M. Stoltze & N. Scharff leg. (ZMUC).

Type locality

TANZANIA: Morogoro Region, Udzungwa Mts, Mwanihana FR, 1800-1850 m asl, montane rain forest.

Colouration

After 31-33 years in alcohol faded, only some darker markings on head and collum left. No traces of a

mid-dorsal stripe.

Distribution and habitat

Known only from Mwanihana FR. Altitudinal range: 1650-1850 m asl. Habitat: montane rain forest.

Coexisting species

Aquattuor submajor Enghoff sp. nov. and Chaleponcus mwanihanensis Enghoff, 2014 occur in the same

area, the former, however, only at lower altitudes.

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Fig. 6. Aquattuor major Enghoff sp. nov., paratype. A. Left gonopod, anterior view. B. Left gonopod,

posterior view. C-D. Telomere tip, two different views, li = lateral incision; pa = palette; mpl = mesal

posterior lamella of telomere; pn = posttorsal narrowing; tt = triangular tooth of telomere. Scale bars:

A-B = 0.1 mm ; C-D = 0.01 mm

European Journal of Taxonomy 150: 1-25 (2015)

Aquattuor longipala Enghoff sp. nov.

urn:lsid:zoobank.org:act:69F309C5-A967-4A68-9776-7C03E879C06D

Fig. 7

Diagnosis

Diameter 1.78-1.90 mm, 50-52 podous rings. Gonopods: lateral incision of coxa (//) deep with rounded

bottom, curving distad, basally delimited by triangular tooth; apical palette (pa ) obliquely rounded, much

longer than broad; telomere describing a c. 270° curve in almost one plane; mesal-posterior lamella of

telomere (mpl) moderately developed; anterior margin of telomere without a triangular tooth; telomere

tip with densely microspiculate margins and internal surface.

Fig. 7. Aquattuor longipala Enghoff sp. nov., paratype. A. Gonopods, anterior view. B. Gonopods,

posterior view. C. Gonopods, oblique apical (central) view. D. Detail of telomere tip. li = lateral incision;

pa = palette; mpl = mesal-posterior lamella of telomere. Scale bars: A-C = 0.1 mm; D = 0.01 mm.

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Etymology

The name is a Latin noun, here in apposition, composed of longi- (long) and pala (palette), and refers to

the shape of the apical part of the gonopod coxa.

Material studied (total: 3 SS)

Holotype

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, West Kilombero Scarp FR,

07°50 , 38.4” S, 36°22 , 17.6” E, montane forest, 1390-1410 m asl, plot Paradiso, 18 Nov. 2000, Frontier

Tanzania leg. (ZMUC).

Paratypes

TANZANIA: 2 $$, same data as holotype (ZMUC).

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, West Kilombero Scarp FR, 07°50 , 38.4” S,

36 0 22’17.6” E, montane forest, 1390-1410 m asl.

Colouration

After 15 years in alcohol still with a clear mid-dorsal, light longitudinal band, flanked by darker areas.

Head and collum with some dark markings.

Distribution and habitat

Known only from West Kilombero Scarp FR. Altitudinal range: 1390-1410 m. Habitat: montane forest.

Coexisting species

Aquattuor udzungwensis Enghoff sp. nov., Chaleponcus netus Enghoff, 2014, and C. circumvallatus

Enghoff, 2014 were found in the same sample as A. longipala Enghoff sp. nov. In addition, C. basiliscus

Enghoff, 2014, C. gracilior Enghoff, 2014, C. ibis Enghoff, 2014, and C. tintin Enghoff, 2014 were

found in West Kilombero Scarp FR.

Aquattuor udzungwensis Enghoff sp. nov.

urn:lsid:zoobank.org:act:60F38BFF-E5EA-4BB0-A691-263DB097502B

Figs 3, 8

Diagnosis

Diameter 1.44-1.69 mm, 44-50 podous rings. Gonopods: lateral incision of coxa (//) deep with rounded

bottom, curving distad; apical palette (pa) rounded-triangular, not longer than broad; telomere describing

a c. 270° curve in almost one plane; mesal-posterior lamella of telomere (mpl) high, basally set off

by 45-90° angle; anterior margin of telomere without a triangular tooth; telomere tip with smooth to

microspiculate margins, internal surface microspiculate.

Etymology

The name is a Latin adjective derived from the Udzungwa Mts, home of this (and several other)

Aquattuor species.

Material studied (total: 21 <$<$)

Holotype

TANZANIA: S, Udzungwa Mts, Iringa Region, Udzungwa Scarp FR, above Chita village, 750 m asl,

23 Oct. -14 Nov. 1984, lowland rain forest, N. Scharff leg. (ZMUC).

European Journal of Taxonomy 150: 1-25 (2015)

Fig. 8. Aquattuor udzungwensis Enghoff sp. nov. A-B, D. Paratype. C. Specimen from West Kilombero

Scarp FR. A. Gonopods in situ. B. Telopodite, ventral view. C. Right gonopod, anterior view.

D. Telomere tip. li = lateral incision; pa = palette; mpl = mesal-posterior lamella of telomere. Scale bars:

A-C = 0.1 mm; D = 0.05 mm

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Paratypes

TANZANIA: 4 SS, same data as holotype, but 25-29 Oct. 1984, pitfall traps (ZMUC).

Referred non-type material

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, West Kilombero Scarp FR, montane

forest, Frontier Tanzania coll.: 1 07°53’19.5” S, 36°23 , 11.6” E, 1100-1300 m asl. Plot Ukami, 30

Nov. 2000 (ZMUC); 6 SS, 07°53’19.5” S, 36°23’11.6” E, 1111 m asl, 23 Nov. 2000 (ZMUC); 7

07°53’19.5” S, 36°23’ 11.6”E, 1126m asl, plot 17-Ukami, 25 Nov. 2000 (ZMUC); 1 $, 07°50 , 38.4” S,

36°22’17.6” E, 1390 m asl, 13 Nov. 2000 (ZMUC); 1 S, 07°50 , 38.4” S, 36°22’17.6” E, 1390-1410 m

asl. Plot Paradiso, 18 Nov. 2000 (ZMUC).

Type locality

TANZANIA: Udzungwa Mts, Iringa Region, Udzungwa Scarp FR, above Chita village, 750 m asl,

lowland rain forest.

Colouration

After 15-31 years in alcohol faded, pale yellowish, head and especially collum contrastingly darker.

Traces of paramedian, dark bands flanking a middorsal, pale stripe.

Distribution and habitat

Known from Udzungwa Scarp FR and from West Kilombero Scarp FR. Altitudinal range: 750-1410 m.

Habitat: lowland rain forest and montane rain forest.

Coexisting species

In Udzungwa Scarp FR, Aquattuor stereosathe Enghoff sp. nov., Chaleponcus circumvallatus Enghoff,

2014, C. hamerae Enghoff, 2014, and C. nikolajscharffi Enghoff, 2014 have been found. In West

Kilombero Scarp FR, Aquattuor longipala Enghoff sp. nov., Chaleponcus netus Enghoff, 2014, and C.

circumvallatus Enghoff, 2014 were found in the same sample as A. udzungwensis Enghoff sp. nov. In

addition, C. basiliscus Enghoff, 2014, C. gracilior Enghoff, 2014, C. ibis Enghoff, 2014, and C. tintin

Enghoff, 2014 were found in West Kilombero Scarp FR.

Aquattuor stereosathe Enghoff sp. nov.

urn:lsid:zoobank.org:act:15CD343E-ABFB-400A-89A0-B25B26E2344C

Figs 3, 9

Diagnosis

Diameter 1.68-1.75 mm, 46-51 podous rings. Gonopods: lateral incision of coxa (//) broad, shallow,

with rounded bottom, basally delimited by triangular tooth; apical palette (pa) rounded, not longer than

broad; telomere describing a 360° curve in three dimensions; mesal-posterior lamella of telomere (mpl)

low; anterior margin of telomere without a triangular tooth; telomere tip with coarsely laciniate margins,

internal surface smooth.

Etymology

The name is a Greek noun, here in apposition, composed of stereo-, which refers to the three-

dimensionality of the curvature of the telomere, and sathe, literally meaning “penis”, here referring to

the telomere.

European Journal of Taxonomy 150: 1-25 (2015)

Material studied (total: 8 ff)

Holotype

TANZANIA: S, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village, 1500 m asl,

2-13 Nov. 1984, N. Scharff leg. (ZMUC).

Fig. 9. Aquattuor stereosathe Enghoff sp. nov., paratype. A. Gonopods, anterior view. B. Gonopods,

oblique lateral view. C. Tip of telomere. D. Gonopods, posterior view, li = lateral incision; pa = palette.

Scale bars: A-C = 0.2 mm; D = 0.01 mm

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Paratypes

TANZANIA: 3 $$, same data as holotype, but 1050 m asl, 26-29 Oct.1984, pitfall traps in intermediate

rain forest (ZMUC); 1 S, same data, but 1400 m asl, 4-5 Nov. 1984 (ZMUC); 3 Iringa Region,

Udzungwa Mts, Udzungwa Scarp FR, Luhega Forest, 08°23’ S, 35°59’ E, 900 m asl, 15 Jul.1990,

J. Fovett, E. Mulungu & F. Hansen leg.

Type locality

TANZANIA: Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village, 1500 m asl.

Colouration

After 25-31 years in alcohol still with a clear mid-dorsal, light longitudinal band, flanked by darker

areas. Head and collum with some dark markings.

Distribution and habitat

Known only from Udzungwa Scarp FR. Altitudinal range: 900-1500 m. Habitat: (intermediate) rain

forest.

Coexisting species

In Udzungwa Scarp FR, Aquattuor udzungwensis Enghoff sp. nov., Chaleponcus circumvallatus

Enghoff, 2014, C. hamerae Enghoff, 2014, and C. nikolajscharffi Enghoff, 2014 have been found.

Species from areas other than the Udzungwa Mountains

In addition to the Udzungwa Mts, species of Aquattuor have been collected in the East Usambara,

Nguru and Rubeho Mts, which all belong to the Eastern Arc Mountains (Burgess et al. 2007), and on

Mt. Kilimanjaro, a much younger, volcanic mountain (Nonnotte et al. 2008).

Aquattuor claudiahempae sp. nov.

urn:lsid:zoobank.org:act:66387356-4B9A-467E-A3CA-4BC76EF8FCAE

Figs 1, 10

Diagnosis

Diameter 1.41-1.58 mm, 50-53 podous rings. Gonopods: lateral incision of coxa (//) poorly developed;

apical palette (pa) subquadratic, with well-developed meso-basal lobe (mbl); telomere describing a

c. 270° curve in almost one plane; mesal-posterior lamella of telomere (mpl) low; anterior margin of

telomere without a triangular tooth; telomere tip densely microspiculate, margin microserrate.

Etymology

The name is a feminine noun in the genitive, referring to Claudia Hemp, Wurzburg, project leader of the

DFG research group Kilimanjaro, in acknowledgement of her great support.

Material studied (total: 26 SS)

Holotype

TANZANIA: S, Kilimanjaro Region, Hai District, Mt. Kilimanjaro, 03°16’07.05” S, 37°18 , 28.68” E,

homegarden plot 2, 1169 m asl, 26 Nov. 2013, S.B. Frederiksen leg. (ZMUC).

Paratypes

TANZANIA: Kilimanjaro Region, Mt. Kilimanjaro: 8 same data as holotype (ZMUC); 1 same

data, but 21 Feb. 2014 (ZMUC); 13 SS, Hai District, 03°13’59.37” S, 37°16 , 09.28”E, coffee plantation

European Journal of Taxonomy 150: 1-25 (2015)

Fig. 10. Aquattuor claudiahempae sp. nov., paratype. A. Gonopods, anterior view. B. Right palette,

showing well developed meso-basal lobe. C. Telomere tip. D. Margin of telomere tip. E. Gonopods,

poterior view, mbl = mesobasal lobe of palette; pa = palette; st9 = sternum of rudimentary 9 th leg-pair.

Scale bars: A, E = 0.1 mm ; B = 0.05 mm ; C = 0.02 mm ; D = 0.01 mm

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

plot 2, 1345 m asl, 20 Nov. 2013, S.B. Frederiksen leg. (ZMUC); 1 Hai District, 03°14 , 34.66” S,

37°15 , 03.91” E, coffee plantation plot 2, 1306 m asl, 11 Mar. 2014, S.B. Frederiksen leg. (ZMUC); 1

same data, but24Feb. 2011, J.Roderleg. (ZMUC); l(?,RomboDistrict, 03°17’57.57”S,37°36’58.59”E,

grassland plot 5, 1303 m asl, 16 Nov. 2011, J. Roder leg. (ZMUC).

Type locality

TANZANIA: Kilimanjaro Region, Hai District, Mt. Kilimanjaro, 03°16’07.05” S, 37 0 18 , 28.68” E,

1169 m asl.

Colouration

Mostly brown, darker on the dorsal side, with a clear mid-dorsal, yellow longitudinal band, lighter

yellowish on the ventral side; legs, head, collum and telson mostly brown (Fig. 1).

Distribution and habitat

Known only from Mt. Kilimanjaro in habitats disturbed by human activities: chagga homegardens (Soini

2005) and coffee plantations. The grassland plot where one specimen was collected is used for fodder

harvest and is thus also under human influence. Altitudinal range: 1169-1345 m asl.

Aquattuor off. claudiahempae sp. nov.

Material studied (total: 1 c?)

TANZANIA: 1 S, Morogoro Region, Kilosa District, Rubeho Mts, 5 km SW of Madizini, 900 m asl,

10-24 Sep. 1983, M. Andersen leg. (ZMUC).

Remarks

Diameter 1.68 mm, 49 podous rings. Colouration faded. Apart from its larger diameter, this specimen

looks completely like A. claudiahempae sp. nov., despite the 350 km distance between the two sites

(cf Discussion).

Aquattuor denticulatus Frederiksen, 2013

Fig. 11

Diagnosis

Diameter 1.58-1.77 mm, 50-52 podous rings. Gonopods: lateral incision of coxa (//) deep, basally

delimited by triangular tooth; apical palette (pa ) rounded, not longer than broad; telomere bending

90° distal after basal lamella, tip of telomere curving laterad (towards coxa); mesal-posterior lamella

of telomere (mpl) low; anterior margin of telomere without a triangular tooth; telomere tip densely

microspiculate.

Etymology

The name refers to the microspiculate (denticulate) telomeral tip (Frederiksen 2013).

Material studied (total: 5 <$<$)

Holotype

TANZANIA: c?. East Usambara Mts, Amani, 1000 m asl, 20 Jul.1980, M. Stoltze & N. Scharff leg.

(ZMUC 00020529).

Paratypes

TANZANIA: East Usambara Mts, Amani: 1 S, 500 m asl, 7 Feb. 1977, H. Enghoff, O. Lomholdt

& O. Martin leg. (ZMUC 00020528); 1 1000 m asl, 27 Jan. 1977, H. Enghoff, O. Lomholdt & O.

European Journal of Taxonomy 150: 1-25 (2015)

Fig. 11. Aquattuor denticulatus Frederiksen, 2013, paratype. A. Gonopods, posterior view. B. Telomere,

distal part, lateral view. C. Gonopods, anterior view. D. Telomere tip. li = lateral incision; bs = basomeral

spine; pa = palette; mpl = mesal-posterior lamella of telomere. Scale bars: A, C = 0.1 mm; B = 0.02 mm;

D = 0.01 mm

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

Martin leg. (ZMUC 00020530); 1 8, 1000 m asl, 15 Jul. 1979, M. Stoltze leg. (ZMUC 00021429); 1 8,

1000 m asl, 28 Dec. 1975, E. Wederkinch leg. (ZMUC 00020610).

Type locality

TANZANIA: East Usambara Mts, Amani, 1000 m asl.

Colouration

After 35—40 years in alcohol, colouration faded, yellowish brown with no clear markings.

Distribution and habitat

Known only from the East Usambara Mts. Altitudinal range: 500-1000 m asl. Habitat: lower montane

forest.

Aquattuor sp.

Material studied (total: 1 5)

TANZANIA: 1 §, Nguru Mts, Mhonda Mission at Turiani, Oct. 1992, M. Andersen leg. (ZMUC).

Remarks

The characteristic limbus places this specimen in Aquattuor , but in the absence of a male, the identity

of the species remains obscure. The Nguru Mts are located roughly midway between the Udzungwa and

the Usambara Mts.

Key to species of Aquattuor Frederiksen, 2013

The key is based on adult males. The diameter and number of podous rings are not absolute characters -

newly collected specimens may fall outside the ranges (and because of the positive correlation between

number of rings and diameter, if a specimens falls outside in one parameter, it is likely to fall outside in

the other as well). Differences between species are subtle, and specimens should be carefully compared

with the diagnoses and illustrations. Specimens collected in new localities are likely to represent new

species.

1. Palette (pa) of gonopods elongate, much longer than broad (Figs 6A, 7A); 50+ podous rings,

diameter >1.7 mm .2

- Palette as long as broad or slightly longer (Figs 5A, 8A, 9A, 10A, 11 A); diameter <1.9 mm.3

2. 53-54 podous rings, diameter >1.9 mm; anterior margin of telomere with a triangular tooth

(Fig. 6A, tt) . A. major Enghoff sp. nov. (Udzungwa: Mwanihana)

50-52 podous rings, diameter <1.9 mm; anterior margin of telomere without triangular tooth

. A. longipala Enghoff sp. nov. (Udzungwa: W Kilombero Scarp)

3. Subdistal part of telomere almost straight, parallel to coxite, telomere tip bent abruptly lateral

(Fig. 11); 50-52 podous rings, diameter 1.6-1.8 mm.

. A. denticulatus Frederiksen, 2013 (E Usambara)

- Telomere describing a smoother curve (Figs 5A, D, 8B-C, 9C, 10E).4

4. Telomere describing a c. 360° curve, curving in three dimensions; telomere tip with serrate margins,

but surface not microspiculate. A. stereosathe Enghoff sp. nov. (Udzungwa: Udzungwa Scarp)

Telomere describing a c. 270°curve, curving almost in one plane only; telomere tip with micro¬

spiculate surface.5

European Journal of Taxonomy 150: 1-25 (2015)

5. Lateral incision (//) of gonopod coxa deep, apical palette (pa) rounded-triangular (Fig. 8A);

<50 podous rings, diameter < 1.7 mm .

. A. udzungwensis Enghoff sp. nov. (Udzungwa: Udzungwa Scarp, W Kilombero Scarp)

- Lateral incision (li) indistinct.6

6. Palette (pa) of gonopods subquadratic, with well-developed meso-basal lobe (mbl) (Fig. 10A-B);

50-53 podous rings, diameter 1.4-1.6 mm. A. claudiahempae sp. nov. (Mt. Kilimanjaro)

Palette more rounded, meso-basal lobe small (Fig. 5A-B); 45—48 podous rings, diameter

1.5-1.9 mm . A. submajor Enghoff sp. nov. (Udzungwa: Mwanihana)

Discussion

Although by far most of the available material of Aquattuor species is from the Udzungwa Mts, the

genus has also been found in the East Usambara, Nguru and Rubeho Mts, all part of the Eastern Arc

Other forested mountains

Kilimanjaro

East Usambara

Rubeho

Udzungwa

Fig. 12. Map of part of Tanzania showing the mountains where Aquattuor species have been found.

Based on Burgess et al. (2007: fig. 1).

ENGHOFF H. & FREDERIKSEN S.B., Aquattuor millipedes from Tanzania

(Figs 12-13), and it is quite likely that Aquattuor will be discovered elsewhere in this ancient mountain

system. The occurrence of A. claudiahempae sp. nov. on Mt. Kilimanjaro appears to be an anomaly,

and three lines of evidence suggest that this species may have been introduced into Mt. Kilimanjaro:

1) it occurs only in disturbed habitats, 2) one specimen which is virtually identical to A. claudiahempae

sp. nov. has been found in the Rubeho Mts, 350 km from Mt. Kilimanjaro, and 3) A. claudiahempae sp.

nov. was not collected during earlier explorations of Mt. Kilimanjaro, including the rather large-scale

Sjostedt Expedition (Attems 1909).

Acknowledgements

SBF was sponsored by the Research-Unit 1246 ( https://www.kilimanjaro.biozentrum.uni-wuerzburg.

de/) of the German Research Foundation (DFG). We are particularly grateful to Claudia Hemp for

making this possible. Research permits were granted by COSTECH, TAWIRI and TANAP (permits nos

2011-346-ER-96-44 and 2014-305-ER-96-44). This work would not have been possible at all without

the impressive collections made by Frontier Tanzania. Our sincere thanks are due to this NGO, as well

as to Nikolaj Scharff and other colleagues at the Natural History Museum of Denmark, and to Juliane

Roder, University of Marburg, who all collected important material and provided various advice, as well

as to Sofia Reboleira, Natural History Museum of Denmark, who provided the photograph in Fig. 1.

MIM ML

IUIM-A

W Kilombero

Scarp

Udzungwa

Scarp

2 Li km

Wooded £i'ii^liiri(l

Woodland

| Open ILiicsl

| Closed forest

O Towns

\l;ijni' mjiiit

9^ Risers

Fig. 13. Map of the Udzungwa Mts showing the forest reserves where Aquattuor species have been

found, and the epithets of the species found. Based on Marshall et al. (2010: fig. 1).

European Journal of Taxonomy 150: 1-25 (2015)

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Manuscript received: 20 August 2015

Manuscript accepted: 8 October 2015

Published on: 6 November 2015

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, Denmark.