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http://dx.doi.org/10.5852/ejt.2016.215

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Research article

urn:lsid:zoobank.org:pub:419BAAE0-E924-41CA-B00C-79FFB5C7E294

A mountain of millipedes IV: Species of Prionopetuhim Attems, 1909,

from the Udzungwa Mountains, Tanzania. With notes on

“P.” fasciatum (Attems, 1896) and a revised species key

(Diplopoda, Spirostreptida, Odontopygidae)

Henrik ENGHOFF

Natural History Museum of Denmark, University of Copenhagen, Universitetsparlcen 15,

DK-2100 Copenhagen 0, Denmark. Email: [email protected]

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

Abstract. Two species of the genus Prionopetalum Attems, 1909, are recorded from the Udzungwa

Mountains: P. asperginis sp. nov. and P. kraepelini (Attems, 1896). Prionopetalum stuhlmanni Attems,

1914, is synonymized under P. kraepelini. Odontopyge fasciata Attems, 1896, is transferred from

Prionopetalum to Aquattuor Frederiksen, 2013, and new illustrations are given. Anew illustrated key to

species of Prionopetalum is provided.

Keywords. Eastern Arc, taxonomy, new species.

Enghoff H. 2016. A mountain of millipedes IV: Species of Prionopetalum Attems, 1909, from the Udzungwa

Mountains, Tanzania. With notes on “P.” fasciatum (Attems, 1896) and a revised species key (Diplopoda,

Spirostreptida, Odontopygidae). European Journal of Taxonomy 215: 1-23. http://dx.doi.org/10.5852/eit.2016.215

Introduction

This is the fourth in a series of articles about the millipedes, especially the endemic Afrotropical family

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

and the Udzungwa Mountains see the first article in the series (Enghoff 2014; see also Enghoff &

Frederiksen 2015 and Enghoff 2016).

Unlike many other genera of Odontopygidae, Prionopetalum Attems, 1909, is well-defined and easily

recognized since the monumental work of Kraus (1960). It is also one of the few odontopygid genera

which have been subject of a subsequent comprehensive review (VandenSpiegel & Pierrard 2009). In

both of these treatments, a key to species is included, in German and French, respectively. Considering

that the vast majority of species of Prionopetalum live in East Africa, where English is much more

widely understood than German and French, a key in English is provided here.

Two species of Prionopetalum , one of them new, have been collected in the Udzungwa Mountains and

are (re)described here. Prionopetalum stuhlmanni Attems, 1914, is shown to be the same species as

P. kraepelini (Attems, 1896). Furthermore, Odontopyge fasciata Attems, 1896, which had been included,

with some doubts, in Prionopetalum by previous authors, is shown to belong in the genus Aquattuor

Frederiksen, 2013. With these adjustments, Prionopetalum now contains 23 described species.

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Material and methods

The bulk of the material for this article comes from the zoological collections of the Natural History

Museum of Denmark, University of Copenhagen (ZMUC). These specimens were collected during field

trips by ZMUC staff and students. Additional specimens from other institutions (see below) were also

examined. All specimens are kept in 70% alcohol.

Specimens were examined in alcohol under a stereo microscope. Specimens for scanning electron

microscopy (SEM) were transferred to 96% ethanol, then to acetone, air-dried, mounted on aluminium

stubs or on pieces of flexible aluminium tape and in turn mounted on stubs, coated with platinum-

palladium and studied in a JEOT JSM-6335F scanning electron microscope.

As in previous articles in this series, only adult males are considered. A total of 22 adult males of the

two Udzungwa species were examined. Figure 1 shows the Udzungwa localities where Prionopetalum

specimens were collected.

See Enghoff (2014) for the description standards used.

[RINGA

20 km

Udzungwa

Scarp

MIKUMI

Wooded grassland

Woodland

| Open forest

m Closed forest

O Towns

/ Ma jor roads

Rivers

TANZANIA

Fig. 1 . Map of the Udzungwa Mountains, showing the collecting localities for Prionopetalum

asperginis sp. nov. (yellow dot) at the southern extremity of the Udzungwa Scarp Forest Reserve and for

P. kraepelini (Attems, 1896) (red diamond) at the eastern edge of the Mwanihana Forest Reserve. Based

on fig. 1 in Marshall et al. (2010).

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

Abbreviations for morphological terms used in the descriptions and on illustrations

amp

mbl

mlf

mml

mmp

mof

prl

pts

pxl

slm

tdp

tpp

anterior metaplical process

basomere

efferent groove

internal canal

lateral coxal tubercle

mesobasal lobe of coxal palette

metaplical longitudinal flange

metaplical mesad lobe

distomesal metaplical process

metaplical oblique/horizontal flange

apical palette of coxa

proplical lobe

post-torsal spine

proximal lobe of telomere

rough area on telomere

solenomere

telomeral distal process

telomeral proximal process

Other abbreviations used in the text

asl

NHMW

VMNH

ZMUC

ZMUH

above sea level

Naturhistorisches Museum, Vienna

Virginia Museum of Natural History

Natural History Museum of Denmark (Zoological Museum)

Zoologisches Museum der Universitat Hamburg

Results

Class Diplopoda Blainville-Gervais, 1844

Order Spirostreptida Brandt, 1833

Family Odontopygidae Attems, 1909

Subfamily Archepyginae Manfredi, 1939

Tribe Prionopetalini Hoffman, 1991

Genus Prionopetalum Attems, 1909

Prionopetalum Attems, 1909: 51.

Type species: Prionopetalum serratum Attems, 1909, by original designation.

Unlike many other genera of Odontopygidae, Prionopetalum is well-defined and quite homogeneous,

not only in non-sexual characters, but also in gonopod structure.

Diagnosis

(Modified after Kraus 1960 and VandenSpiegel & Pierrard 2009, excluding some non-gonopodal

characters which were mentioned by these authors but which are of no diagnostic value.)

Prionopetalini in which the anal valves have a raised rim, a well-developed dorsal spine and sometimes

a smaller ventral one; limbus with simple, pointed denticles (true of all species after removal of

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Table 1 . Numbers of podous rings and body diameter of adult males of species of Prionopetalum. One

has been subtracted from published “segment” numbers because these include the telson. None of the

species are known to have apodous rings between the last podous ring and the telson.

No. of podous rings

Diameter (mm)

Source

P. aculeatum

Attems 1914

P. asperginis sp. nov.

60-65

4.4^F9

this study

P. bifidum

68-71

5.6-6.8

VandenSpiegel & Pierrard 2009

P. clarum

“nearly fourty-three”

5.5

Chamberlin 1927

P. cornutum

Kraus 1958

P. coronatum

3.7

Kraus 1958

P. dentigerum

6.0

this study 1

P. etiennei

2.5

this study 2

P. exaratum

2.5

Attems 1938

P. frundsbergi

7.6

this study 3

P. fryeri

Turk 1956

P. glomeratum

2.5

Attems 1935

P. kraepelini

61-65

3.2-3.6

Attems 1896; this study

P. lindi

58-59

3.3-3.8

VandenSpiegel & Pierrard 2009

P. megalacanthum

4.3

Attems 1912

P. ndelei

61-63

4.2

VandenSpiegel & Pierrard 2009

P. pulchellum

3.0

Kraus 1960

P. serratum

62-66

5.6

Attems 1909

P. suave

Attems 1896

P. tanganjikum

Verhoeff 1941

P. tricuspis

5.5

Brolemann 1920

P. urbicolum

61-64

4.5

Carl 1909

P. xerophilum

c. 60

4.5

Carl 1909

1 P. dentigerum. diameter not provided by Verhoeff (1941). Values based on f from Tanzania, Pwani Region,

Kisarawe District, Ruvu South Forest Reserve, 140 m asl, 6°57'27" S, 38°50'51" E, date unknown, leg. Frontier

Tanzania, det. H. Enghoff (ZMUC).

2 P. etienner. diameter not provided by Demange (1982). Values based on S from Guinee-Bissau, Buba, 9-11 Jun.

1989, leg. A. van Harten & M. Neves., det. H. Enghoff (ZMUC).

3 P. fnmdsbergi: no published information. Values based on f from Kenya, Bushwackers near Kibwezi, 30 Dec.

1982, leg. C.C. Kinze et al, det. El. Enghoff (ZMUC).

“P. ” fasciatum , see below). Male legs with ventral soft pads on postfemur and tibia, except on anteriormost

and posteriormost legs (true of all species after removal of “P ” fasciatum , see below).

Gonopods. Coxal metaplica on its basal part with a large longitudinal mesad flange {mlf), separated by

a deep sinus from an oblique-horizontal sub-semicircular mesad flange (mof). Telopodite with a post-

torsal (“femoral”) spine (pts), but without a (“tibial”) spine near the origin of the solenomere, divided

into solenomere and telomere shortly after post-torsal narrowing. Solenomere ( slm ) simple, whip-like,

without outgrowths (exception: P. fryeri (Turk, 1956) with a short accessory branch at c. 2 A of the

ENGHOFF H., Prionopetalum millipedes from the Udzungwa Mts

solenomere’s length). Telomere with two characteristic processes: a variously shaped, often species-

specific proximal process ( tpp ) which projects at ± right angles from the main telomere axis, and a long,

slender distal process (tdp) which is armed with a row of spines or thorns. (In the terminology of Kraus

(1960), tpp is “eine hornartige Spitze oder auch eine entsprechende, schalige Famelle”, and tdp is “der

meist auffallende schlanlcere Distalabschnitt.gezackt oder bezahnt”. For VandenSpiegel & Pierrard

(2009), tpp is a “processus basal du tarse”.)

Body size

Published body diameters for adult males range from 2.5 mm (P. exaratum, P. glomeratum ) to 6.8 mm

(P. bifidum ), but a male of P. frundsbergi in the ZMUC collection has a diameter of 7.6 mm. Published

numbers of podous rings range from 57 (P. exaratum) to 71 (P. bifidum ), but the above-mentioned male

of P. frundsbergi has 72 podous rings. Table 1 and Fig. 2 summarise the size information and give

detailed information for the two Udzungwa species. One has been subtracted from published “segment”

numbers because these have traditionally included the telson. Two records have been omitted: “nearly

fourty-three” for P. clarum (Chamberlin, 1927) and 93 for P. tanganjikum Verhoeff, 1941 - the latter

number is probably a lapsus calami.

"nj 5

■o

QD □ _ □□

a ^ p°

■■

0 _ ♦

♦ P. kraepelini Udzungwa

0 i ♦ O o ♦

O P. kraepelini - Attems 1896

□ P. asperginis sp. nov.

XP. dentigerum

AP. etiennei

o A

#P. frundsbergi

Oother spp.

—1—

-1-

-I-

-1-

Number of podous rings

Fig. 2. Body size (body diameter / number of podous rings) of $$ of Prionopetalum spp. Based on

original measurements and data from the literature (see Table 1). For “other spp.” the entries are median

values of the intervals in Table 1 .The report of “93 (1) Rumpfringen” (corresponding to 92 podous rings)

for P. tanganjikum by Verhoeff (1941) has been omitted, as has Chamberlin’s (1927) “nearly fourty-

three” segments for P. clarum ; both are regarded as quite unlikely and are probably erroneous.

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Included species

P aculeatum Attems, 1914 - Somalia, Kenya

P. asperginis sp. nov. - Tanzania

P. bifidum VandenSpiegel & Pierrard, 2009 - Tanzania

P. clarum (Chamberlin, 1927) -D.R. Congo

P cornutum Kraus, 1958-D.R. Congo

P coronatum Kraus, 1958-D.R. Congo

P. dentigerum Verhoeff, 1941 - Tanzania, Rwanda

P. etienneiDQmmgQ, 1982-Gambia, Guinee-Bissau, Senegal

P exaratum (Attems, 1938)-D.R. Congo

P. frundsbergi (Attems, 1927) - Kenya, Tanzania

P. firyeri (Turk, 1956) - Malawi, Zambia

P. glomeratum Attems, 1935 - D.R. Congo

P. kraepelini ( Attems, 1896) - Tanzania

P. lindi VandenSpiegel & Pierrard, 2009 - Tanzania

P. megalacanthum Attems, 1912 - D.R. Congo, Rwanda

P. ndelei VandenSpiegel & Pierrard, 2009 - Central African Republic

P pule helium Kraus, 1960 - Mozambique

P. serratum Attems, 1909 - Kenya, Tanzania

P. suave (Gerstacker, 1873)-Tanzania

P. tanganjikum Verhoeff, 1941 - Tanzania

P. tricusp is Brolemann, 1920-Kenya

P urbicolum (Carl, 1909) - Tanzania

P. xerophilum (Carl, 1909) - Rwanda, Tanzania

Prionopetalum asperginis sp. nov.

um:lsid:zoobank.org:act:FE9E9A13-4D50-40E9-8B6D-3D40AEA0ElA2

Figs 3-4, 9W, 10D, 11K

Diagnosis

Differs from congeners by the combination of a laterally smooth gonopod coxa, a pointed apical mesad

metaplical process subtended by a rounded mesad lobe, a simple, fingerlike proximal telomere process

and a simple distal telomere process without secondary branches.

Etymology

The species is named after the recently discovered Kihansi spray toad, Nectophrynoides asperginis

Poynton, Howell, Clarke & Lovett, 1999; see “Distribution and habitat”.

Material studied (total: 15 SS)

Holotype

TANZANIA: $ Udzungwa Mts, Kihansi, 8°24' S, 36°21' E, “forest site”, Jun -Aug. 1997, L Zilihona

leg. (ZMUC).

Paratypes

TANZANIA: 12 same data as holotype (ZMUC); 1 $ Iringa Region, Mufindi District, Udzungwa

Scarp Forest Reserve, 8°31.58' S, 35°53.9F E, 750 m asl, 5-12 Mar. 1996, GUV28-2, Proj. S.H.

McKamey et al. leg. (ZMUC); 1 $ Morogoro Region, Kilombero District, Lower Kihansi Project,

Udagaji Gorge, 7 Nov. 1997, Jan Kielland leg. (VMNH).

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

Fig. 3. Prionopetalum asperginis sp. nov., paratype from Kihansi, “forest site”. A-B. Hind end.

A. Fateral view. B. Posterior view. C. Fimbus. D-F. Right gonopod coxa. D. Anterior view (light blue:

mounting tape). E. Mesal view. F. Posterior view. G. Transverse section of right gonopod basomere

(circle: bundle of tracheae). H. Cluster of tracheae next to internal canal. Abbreviations: amp = anterior

metaplical process; eg = efferent groove; ic = internal canal; mlf = metaplical longitudinal flange;

mml = metaplical mesad lobe; mmp = distomesal metaplical process; mof= metaplical oblique/horizontal

flange ;prl = proplical lobe. Scale bars: A, B, D-F = 0.1 mm; C, H = 0.001 mm ; G = 0.01 mm

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Type locality

TANZANIA: UdzungwaMts, Kihansi, 8°24' S, 36°21' E.

Fig. 4. Prionopetalum asperginis sp. nov., paratype from Kihansi, “forest site”, right gonopod telopodite.

A. Anterior and slightly mesal view. B. Posterior view. C. Basal view. D. Apical view. E. Apical-mesal

view. F. Mesal and slightly posterior view. Abbreviations: ba = basomere; pts =post-torsal spine (broken);

pxl = proximal lobe of telomere; ra = rough area of telomere; slm = solenomere; tdp = distal process of

telomere; tpp = proximal process of telomere. Scale bars = 0.1 mm.

ENGHOFF H., Prionopetalum millipedes from the Udzungwa Mts

Description

Size. Fength c. 6!4 cm, diameter 4.4-4. 9 mm, 60-65 podous rings, no apodous rings in front of telson.

Colour. After 18 years in alcohol a lm ost uniform light brown. No lighter dorsal markings.

Anal valves (Fig. 3A-B). Each with a long, pointed dorsal spine, no ventral spine, marginal rim raised,

with 3 setae on very poorly demarcated tubercles.

Fimbus (Fig. 3C). With triangular, almost equilateral, pointed lobes, external surface of lobes densely

striate.

Male legs. Postfemora and tibiae with large, soft pads, except on first four to five and several

posteriormost leg pairs.

Gonopod coxa (Fig. 3D-F). Fateral margin almost straight, entirely smooth. Mesal margin of proplica

straight, proplical lobe (prl) in anterior view almost hidden behind apical expansion of metaplica. Basal

part of metaplica with large longitudinal mesad flange ( mlf ), separated by a deep sinus from an oblique-

horizontal, sub-semicircular mesad flange (moj ), apical part of metaplica expanded anteriad, forming a

pointed process {amp) covering proplical lobe, a rounded mesad lobe ( mml) and a slender disto-mesad

process ( mmp ).

Gonopod telopodite (Fig. 4). A well-developed post-torsal spine (“femoral spine”, pts) just before post-

torsal narrowing (broken on illustrated specimen). Solenomere ( slm ) simple, slender. Telomere with

a large, proximal lobe {pxl ) and a rough area (ra) on apical surface, further distally divided into two

processes. Proximal telomere process (tpp) parallel-sided, apically rounded. Distal telomere process

(tdp) only slightly broader than solenomere, apically with one margin denticulate (Fig. 4F).

Distribution and habitat

Known only from the southern part of the Udzungwa Mts, Udzungwa Scarp Forest Reserve and the

Kihansi area. The latter area has become famous because of the Kihansi spray toad, Nectophrynoides

asperginis Poynton, Howell, Clarke & Fovett, 1999, which occurred in the Kihansi area but is now

regarded as extinct in the wild although a reintroduction programme was started in 2013 (IUCN SSC

Amphibian Specialist Group 2015). Zilihona et al. (1998) described in detail the area where I. Zilihona

collected her specimens. Altitudinal range 550-750 m asl (cf. Zilihona et al. 1998).

Coexisting species

The sample collected by I. Zilihona also contains another, much smaller odontopygid which will be

described in a forthcoming paper.

Remarks

Fig. 3G shows a transverse section (= break) of the basal part (basomere, ba) of the gonopod telopodite.

The picture clearly shows how the canal (eg) that continues to the tip of the solenomere is formed as

a groove near the surface. This canal has been supposed to be a sperm canal (“Samenrinne” of Kraus

1966, e.g., his figs 28-29), but considering that nothing is known about its function, the neutral term

“efferent groove” is preferable (cf. Enghoff 2011). There is also what looks like an entirely internal

canal (ic) which is accompanied by dense bundles of tubuli, probably tracheae (Fig. 3H), similar to those

recently described by Reboleira et al. (2015) from the base of the vulva of a cambalid millipede. The

extent and function of this canal is unknown.

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Prionopetalum kraepelini (Attems, 1896)

Figs 5-6, 9R, 111

Odontopyge kraepelini Attems, 1896: 37.

Prionopetalum stuhlmanni Attems, 1914: 210. New synonymy

Odontopyge pardalis - Attems 1896: 39.

Prionopetalum pardalis - Attems 1909: 52.

Prionopetalum kraepelinii — Attems 1914: 210.

Not Spirostreptus pardalis Gerstacker, 1873.

Diagnosis

Differs from all congeners by the multi-cusped proximal telomere process (Fig. 6F).

Material studied (total: 9 ff)

Syntypes

TANZANIA: If, 1 $, Mhonda (Unguru), 6 Sep. 1888, F. Stuhlmann leg. (ZMUH).

Other material

TANZANIA: 1 f, Tewa (Usambara), 26 Apr. 1888, F. Stuhlmann leg., holotype of Prionopetalum

stuhlmanni (ZMUH); 3 ff, Morogoro Region, Mang’ula, at Udzungwa Ecological Monitoring Centre,

300 m, 7°50'56" S, 36°53T7" E, 28-31 Sep. 2012, T. Pape leg. (ZMUC); 4 ff, Morogoro Region,

Udzungwa Mts, Udzungwa Ecological Monitoring Centre, Mang’ula, 07°50'44.9" S, 36°53'28.2" E,

339 m, 18-20 Jan. 2014, T. Pape & N. Scharff leg. (ZMUC).

Type locality

TANZANIA: Tanga Regon, Lushoto District, Usambara Mts, Lewa.

Description (male)

Size. Length c. 5 cm, diameter 3.2-3.6 mm, 61-65 podous rings, no apodous rings in front of telson.

Colour. After 1-3 years in alcohol head amber below antennae, blackish above; antennae blackish brown;

collum blackish brown with light margins; body rings light brown below ozopores and immediately

above, dorsally blackish brown with broad middorsal light spot; telson and legs medium brown.

Anal valves. Each with a long, pointed dorsal spine and a much smaller ventral one, marginal rim

raised, with 3 setae on very poorly demarcated tubercles.

Limbus (Fig. 5F). With triangular, almost equilateral, pointed lobes, external surface of lobes densely

striate.

Male legs. Postfemora and tibiae with large soft pads, except on first four to five and several posteriormost

leg pairs.

Gonopod coxa (Fig. 5A-E). Lateral margin almost straight, entirely smooth or with a tiny tubercle (It) at

c. 2 A height. Mesal margin of proplica straight, proplical lobe (prl ) in anterior view hidden behind apical

expansion of metaplica. Basal part of metaplica with large, longitudinal mesad flange (ml/), separated

by a deep sinus from an oblique-horizontal, two-lobed mesad flange (mof), apical part of metaplica

expanded anteriad to form a process (amp) covering proplical lobe and produced disto-mesad into

slender, apically bifid process (mmp).

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

Fig. 5. Prionopetalum kraepelini (Attems, 1896), specimen from Mang’ula, 339 m asl. A-C. Right

gonopod coxa. A. Anterior view. B. Anterior-mesal view. C. Mesal view. D-E. Feft gonopod.

D. Posterior view. E. Anterior-mesal view. F. Midbody-dorsal limbus. G. Row of intercalary micro¬

scutes with knobs, from midbody metazona. Abbreviations: amp = anterior metaplical process; It =

lateral coxal tubercle; mlf= metaplical longitudinal flange; mmp = distomesal metaplical process; mof=

metaplical oblique/horizontal flange; prl = proplical lobe. Scale bars: A-E = 0.1 mm; F-G = 0.001 mm.

European Journal of Taxonomy 215 : 1-23 ( 2016 )

Fig. 6. Prionopetalum kraepelini (Attems, 1896), specimen from Mang’ula, 339 m asl. Gonopod

telopodite. A, C-D, F. Right gonopod telopodite. A. Posterior view. C. Apical (ventral) view. D. Anterior

view. F. Telomere distal process. B. Tip of left solenomere. E. Teft gonopod, (posterior-) mesal view.

Abbreviations: pts = post-torsal spine; pxl =proximal lobe of telomere; slm = solenomere; tdp = telomeral

distal process; tpp = telomeral proximal process. Scale bars: A, C-D, E = 0.1 mm; B, F = 0.2 mm.

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

Gonopod telopodite (Figs 5D-E, 6). A well-developed post-torsal spine (“femoral spine”, pts) inserted

just before post-torsal narrowing. Solenomere ( slm ) simple, slender. Telomere with a large, proximal

lobe (pxl). Proximal telomere process (tpp) elaborate, with a longitudinal 3-5-dentate flange and a

slender apical process, in certain views (Fig. 5E) recalling the head profile of cartoon character Woody

Woodpecker. Distal telomere process ( tdp ) only slightly broader than solenomere, apically with one

margin denticulate (Fig. 6F).

Distribution and habitat

In addition to the newly collected material from the Udzungwa Mts, Udzungwa Ecological Monitoring

Centre, Mang’ula ( http://www.udzungwacentre.org/ ) at 300-339 m asl, O. kraepelini has been recorded

from several other sites, all in Tanzania: Morogoro Region, Mhonda (Nguru) (type locality); Tanga

Region, Fushoto Distr. Usambara Mts, Fewa (type locality of P. stuhlmanni ); Arusha Region, Arusha

District, near Fake Babati; Dar es Salaam (the latter two records by VandenSpiegel & Pierrard 2009).

Coexisting species

At Mang’ula, P. kraepelini was collected together with another odontopygid species which will be

described in a forthcoming article.

Remarks

Two of the four species of Odontopyge (re)described by Attems (1896) distinguished themselves by the

possession of a peculiar, multi-cusped process on the gonopod telopodite. This process, labelled c on

Attems’ figures, corresponds to the proximal telomere process (tpp) in the sense of the present paper. The

two species in question were O. kraepelini , described as new, and a species which Attems identified as

O. pardalis (Gerstacker). He later (Attems 1914) realised that this was not the real pardalis Gerstacker

and offered the replacement name Prionopetalum stuhlmanni Attems, 1914 ( pardalis has subsequently

been transferred to the genus Calyptomastix Hoffman & Howell, 2012).

The two species, kraepelini and stuhlmanni , are very similar indeed according to the descriptions and

illustrations offered by Attems (1896). The only substantial apparent difference concerns the profile of

the gonopod coxa. Attems (1896) provided two gonopod drawings of stuhlmanni (as pardalis ), and one

of kraepelini. The three drawings are all quite different regarding the gonopod coxa profiles, but this is

due to the fact that two of them, the one of kraepelini (Attems’ fig. 1) and one of those of stuhlmanni

(Attems’ fig. 8), are based on gonopods macerated in KOH, whereas his fig. 7 (of stuhlmanni ) is based

on unmacerated gonopods. Attems’ fig. 7 is fully compatible with the present illustrations (Figs 5-6)

of specimens from Mang’ula, and side-by-side comparisons of the body and gonopods of Mang’ula

specimens with the holotype of stuhlmanni reveal no differences (see Fig. 2 for agreement in body size).

The male syntype of kraepelini is devoid of its gonopods, and these are not retrievable elsewhere. I

interpret the apparent differences between macerated gonopods of kraepelini ( Attems 1896: fig. 1) and

stuhlmanni (Attems 1896: fig. 8) as being artificial and possibly due to different durations of the KOH

maceration, and I therefore synonymize the two names.

Knob-like intercalary cuticular micro-scutes were observed in this species (Fig. 5G), cf. Enghoff (2014).

Under the name Prionopetalum stuhlmanni this species is a popular pet millipede; see, e.g., Sigling

(2010) and http://www.diplopoda.de/index.php (accessed 29 February 2016).

The status ^/“Prionopetalum” fasciatum (Attems , 1896)

Attems (1896) described Odontopyge fasciata from Zanzibar, but later (1914) transferred the species

to Prionopetalum without further comment. Brolemann (1920) redescribed the species based on new

European Journal of Taxonomy 215: 1-23 (2016)

material from Zanzibar and expressed doubts about the genus-level classification of it, suggesting

that it might belong in a separate subgenus of Prionopetalum. Kraus (1960), however, accepted the

inclusion of fasciata in Prionopetalum. In the most recent treatment of Prionopetalum VandenSpiegel &

Pierrard (2009) discussed fasciata , noticing that it differs from other species of Prionopetalum in several

characters: the limbus is almost straight, not serrate; there are no ventral tibial pads on male legs; and

the proximal telomeral process ( tpp ) is a simple, rounded lobe. The latter authors reported further new

material from Zanzibar as well as from Bagamoyo District, Vula Mountain, Pongwe, on the Tanzanian

mainland.

After having examined several specimens from Zanzibar I can now offer a re-classification of this

species:

Aquattuor fasciatus (Attems, 1896) comb. nov.

Figs 7-8

Odontopyge fasciata Attems, 1896: 40.

Prionopetalum fasciatum - Attems 1914: 210. — Brolemann 1920: 123. — Kraus 1960: 86. —

VandenSpiegel & Pierrard 2009: 152. — Enghoff et al. 2016.

Aquattuor aff. claudiahempae - Enghoff & Frederiksen 2015: 19.

Material studied

Syntypes

TANZANIA: 1 3 (+ one extra set of gonopods), 6 $ $/juvs, Zanzibar, Kibueni (ZMUH).

Other material

TANZANIA: 3 33, Zanzibar, Kizimkazi, at foot of mango tree, 17 Jun. 1979, M. Stoltze leg. (ZMUC);

1 3 , Zanzibar, the sultan’s palace, 16 Jun. 1979, M. Stoltze leg. (ZMUC).

In addition. Dr. Nesrine Akkari has kindly examined syntypes of O. fasciata in NHMW and has placed

information and images at my disposal.

Remarks

The single intact male among the examined syntypes has 53 podous rings (no apodous rings in front

of the telson) and a diameter of 1.91 mm. A syntype at NHMW has the same number of rings and a

diameter of 1.86 mm The specimens from Kizimkazi are all broken, the males have maximum body

diameters of 1.55-1.62 mm. The male from the sultan’s palace has 51 podous rings (no apodous rings in

front of the telson) and a diameter of 1.86 mm.

The limbus (Fig. 7A) consists of large, rectangular, easily detached flaps, characteristic of the genus

Aquattuor.

The gonopods (Figs 7B-F, 8) are indistinguishable from those of Aquattuor claudiahempae Enghoff &

Frederiksen, 2015 (cf. their fig. 15). Noticeable special similarities include the profile of the apical

palette (pa), the well-developed mesobasal lobe ( mbl ) of the coxal palette, and the partly micro-serrate

margin of the telomere (Fig. 7 E, arrow).

The characteristic limbus was not noted by previous authors. Attems (1896) and Brolemann (1920) did

not mention the limbus at all. VandenSpiegel & Pierrard (2009: fig. lb) illustrated the limbus, but their

SEM micrograph only showed an irregularly wavy margin. The large limbus lobes were probably all

broken off in the specimens examined by these authors (compare the left part of Fig. 7A with fig lb

ENGHOFF FL, Prionopetalum millipedes from the Udzungwa Mts

Fig. l.Aquattuor fasciatus (Attems, 1896) comb, nov., specimen from Zanzibar, Kirimlcazi. A. Fimbus.

B-C. Feft gonopod. B. Anterior view. C. Posterior view. D-F. Feft telomere. D. Tip, basal-anterior

view. E. Subdistal part, posterior view (bold arrow points to a microserrate lobe on the margin). F. Tip,

apical-posterior view. Abbreviations: mbl = mesobasal lobe of coxal palette; pa = apical palette of coxa.

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

European Journal of Taxonomy 215: 1-23 (2016)

in VandenSpiegel & Pierrard 2009). In the syntypes from ZMUH the limbus seems to be completely

worn off. However, Dr. Nesrine Akkari has kindly provided images based on the syntypes of O. fasciata

in NHMW which clearly show the Aquattuor type of limbus, as well as confirm the great similarity

in gonopod structure between this species and A. claudiahempae (Fig. 8). The gonopods of the intact

male syntype (now dissected by me) closely agree with the illustrations in Figs 7-8. The extra set of

gonopods among the syntypes very much resembles fig. 6 in Attems (1896), which is based on macerated

gonopods, and they were most probably the model for this drawing.

The syntypes of fasciatus as well as the Zanzibar specimens collected by M. Stoltze are very similar to

A. claudiahempae Enghoff & Frederiksen (2015), and it is possible that the latter name should be regarded

as a junior synonym of A. fasciatus. However, the body diameter of the specimens from Kizimkazi

is at the high end, and partly larger, compared to A. claudiahempae from Mt. Kilimanjaro. The two

intact specimens from Zanzibar examined here, viz., the intact syntype and the male from the sultan’s

palace, are definitely much larger. The latter specimens are more similar in size to A. cf. claudiahempae ,

recorded from the Morogoro Region, Kilosa District, Rubeho Mts by Enghoff & Frederiksen (2015).

Attems (1896) gave the (horizontal) diameter of A. fasciatus as 2 mm. Brolemann (1920) gave 55

podous rings (= “56 segments ... 1 segment apode”) and a diameter of 1.70 mm for an adult male.

Whereas “2 mm ” is much more than any other specimen in this complex, Brolemann’s values would fit

A. claudiahempae by extrapolation.

The situation is obviously complicated, and for the time being, A. fasciatus and A. claudiahempae are

kept as separate species.

A key to the species of Prionopetalum

This key builds extensively on previous keys provided by Kraus (1960) and VandenSpiegel & Pierrard

(2009). The new species described here is included, but Aquattuorfasciatus is excluded, andP stuhlmanni

is treated as a synonym of P. kraepelini (cf. above). I follow VandenSpiegel & Pierrard (2009) in not

Fig. 8. Aquattuor fasciatus (Attems, 1896) comb, nov., syntype of Odontopyge fasciata (NHMW 2672).

A. Gonopods, anterior view. B. Left gonopod, posterior view. Scale bars = 0.5 mm. Photographs:

N. Akkari.

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

considering Spinotarsus werneri Attems, 1910 as a species of Prionopetalum , although Kraus (1960)

suggested such a relationship (but still listed S. werneri under ’’species incertae sedis ”).

1. Coxa with a straight, mostly long lateral spine pointing basad, laterobasad or laterad (Fig. 9A-F)

- Coxa different, sometimes with another type of lateral process, but not a straight basad, laterobasad

or laterad spine.13

2. Fateral coxal spine directed laterad, set off by right angles from main coxal margin (Fig. 9A).

. P. megalacanthum Attems, 1912

- Fateral coxal spine different.3

3. Fateral coxal spine directed basad, in part overlying main body of coxa (Fig. 9B).

. P. frundsbergi (Attems, 1927)

- Fateral coxal spine different.4

4. Tip of distal telomere process ( tdp ) divided into two equal branches (Fig. 10A). Gonopod coxa

profile as Fig. 9C. P. bifidum VandenSpiegel & Pierrard, 2009

- Tip of distal telomere process not divided into two equal branches, but one or two small subapical

processes may be present.5

5. Distal telomere process (tdp) ending in three dark spines (Fig. 10B). Gonopod coxa profile as

Fig. 9D . P. tricuspis Brolemann, 1920

- Tip of distal telomere process at most with a single accessory process/spine .6

6. Proximal telomere process ( tpp ) distally expanded, asymmetrically club-shaped (Fig. 11A-F).7

- Proximal telomere process not strongly expanded distally (Fig. 11G-K).12

7. Distal telomere process (tdp) with a small accessory process (Fig. 10C) .8

- Distal telomere process without accessory process (like Fig. 10D) .9

8. Fateral coxal spine relatively short, directed almost basad (Fig. 9E). Proximal telomere process

(tpp) moderately asymmetrical (Fig. 11 A). P. aculeatum Attems, 1914

- Fateral coxal spine directed almost laterad (Fig. 8F). Proximal telomere process strongly asym¬

metrical (Fig. 11B). P. ndelei VandenSpiegel & Pierrard, 2009

9. Coxa mesapically rounded (Fig. 9G). Proximal telomere process as in Fig. 11C.

. P. serratum Attems, 1909

- Coxa mesapically with a spinous projection (Fig. 9H-J, arrows) .10

10. Distal margins of proximal telomere process (tpp) smooth (Fig. 10D). Gonopod coxa profile as

Fig. 9H. P. dentigerum Verhoeff, 1941

- Distal margin of proximal telomere process wavy or with a denticle (Fig. 11E-F) .11

11. Distal margin of proximal telomere process (tpp) wavy (Fig. 11E). Gonopod coxa profile as Fig. 91

. P. tanganjikum Verhoeff, 1941

- Distal margin of proximal telomere process with a single denticle (Fig. 1 IF, arrow). Gonopod

coxa profile as Fig. 9J. P. xerophilum (Carl, 1909)

European Journal of Taxonomy 215: 1-23 (2016)

Fig. 9. Prionopetalum spp., outlines of distal part of gonopod coxa. The species appear in the sequence

in which they key out in the identification key. A. P. megalacanthum (after a specimen in NHMW).

B. P. frundsbergi (based on Kraus 1960). C. P. bifidum (based on VandenSpiegel & Pierrard 2009).

D. P. tricuspis (based on Brolemann 1920). E. P. aculeatum (based on Kraus 1960). F. P. ndelei

(based on VandenSpiegel & Pierrard 2009). G. P. serratum (based on Kraus 1960). H. P. dentigerum

(based on Kraus 1960). I. P. tanganjikum (based on Kraus 1960). J. P. xerophilum (based on Kraus

1960). K. P. clarum (based on Kraus 1960). L. P. pulchellum (based on Kraus 1960). M. P. etiennei

(based on Demange 1982). N. P. lindi (based on VandenSpiegel & Pierrard 2009). O. P. coronatum

(based on Kraus 1958). P. P. exaratum (based on Kraus 1960). Q. P. cornutum (based on Kraus 1960).

R. P. kraepelini (orig.). S. P. glomeratum (based on Attems 1935). T. P. urbicolum (based on Kraus

1960). U. P. suave (based on Kraus 1960). V. P. fryeri (based on Kraus 1960). W. P. asperginis sp. nov.

(orig). Not to scale.

ENGHOFF H., Prionopetalum millipedes from the Udzungwa Mts

12. Proximal telomere process (tpp) straight, apically rounded (Fig. 11G). Body diameter 5.5 mm.

Gonopod coxa profile as Fig. 9K. P. clarum (Chamberlin, 1927)

- Proximal telomere process slightly curved, pointed (Fig. 11H). Body diameter 3 mm. Gonopod

coxa profile as Fig. 9F. P. pulchellum Kraus, 1960

13. Coxa with a strongly curved lateral spine (Fig. 9M-N) .14

- Fateral coxal spine, if present, not strongly curved .15

14. Gonopod coxa profile as Fig. 9M. Body diameter 2.5 mm . P. etiennei Damange, 1982

- Gonopod coxa profile as Fig. 9N. Body diameter 3.3-3.8 mm.

. P. lindi VandenSpiegel & Pierrard, 2009

15. Fateral coxal spine sizeable, directed apicad and partly overlying main body of coxa. Coxa

apically bifid (Fig. 90-Q).16

- Coxa at most with a tiny lateral spine or bump (Fig. 9R-W) .18

16. Both apical branches of coxa pointed, the mesal branch hook-like (Fig. 90) .

. P. coronatum Kraus, 1958

- Mesal-apical branch of coxa not pointed (Fig. 9P-Q) .17

17. Gonopod coxa profile as Fig. 9P. P. exaratum (Attems, 1938)

- Gonopod coxa profile as Fig. 9Q . P. cornutum Kraus, 1958

18. Proximal telomere process (tpp) with several lobes along one side (Fig. 111). Gonopod coxa

profile as Fig. 9R. P. kraepelini (Attems, 1896)

- Proximal telomere process at most with a single lateral lobe (Fig. 11J-K) .19

19. Coxal apex hoolc-like (Fig. 9S). Body diameter 2.5 mm. P. glomeratum Attems, 1935

- Coxal apex not hoolc-like (Fig. 9T-W). Body diameter 4.4-5 mm .20

20. Coxal apex with three mesad processes (Fig. 9T) . P. urbicolum (Carl, 1909)

- Coxal apex with two mesad processes (Fig. 9U-W) .21

Fig. 10. Prionopetalum spp., outlines of distal telomere process (tdp). A. P bifidnm (based on

VandenSpiegel & Pierrard 2009). B. P. tricuspis (based on VandenSpiegel & Pierrard 2009).

C. P aculeatum (based on Attems 1914). D. P. asperginis sp. nov. (orig.). Not to scale.

European Journal of Taxonomy 215: 1-23 (2016)

21. Both mesad processes of coxal apex pointed (Fig. 9U). P. suave (Gerstacker, 1873)

- Subapical mesad process of coxal apex rounded (Fig. 9V-W).22

22. Gonopod coxa profile as Fig. 8V. Solenomere with a short accessory branch at c. 2 A of its length.

Proximal telomere process as Fig. 11J. P. fryeri (Turk, 1956)

- Gonopod coxa profile as Fig. 8W. Solenomere without an accessory branch. Proximal telomere

process as Fig. 11K. P. asperginis sp. nov.

Fig. 11. Prionopetalum spp., outlines of proximal telomere process (tpp) . A. P. aculeatum (based

on Attems 1914). B. P. ndelei (based on VandenSpiegel & Pierrard 2009). C. P. serratum (based on

Attems 1909). D. P. dentigerum (based on Kraus 1960). E. P. tanganjikum (based on Kraus 1960).

F. P. xerophilum (based on Kraus 1960). G. P. clarum (based on Kraus 1960). H. P. pulchellum (based

on Kraus 1960). I. P. kraepelini (orig.). J. P. fryeri (based on Kraus 1960). K. P. asperginis sp. nov.

(orig.). Not to scale.

ENGHOFF Ft., Prionopetalum millipedes from the Udzungwa Mts

Discussion

Among the Odontopygidae from the Udzungwa Mts studied so far, species of Prionopetalum occur

at the lowest altitudes: 300-339 m asl for P. kraepelini and 550-750 m asl for P. asperginis sp. nov.

In comparison, the altitudinal span of the Udzungwa endemic Chaleponcus dabagaensis group is

1390-2100 m asl (Enghoff 2014), of Udzungwa Aquattuor species 750-1800 (1850) m asl (Enghoff &

Frederiksen 2015), and of Udungwa Geotypodon species 1145-1500 m asl (Enghoff 2016). This is in

line with the general tendency of Prionopetalum species to occur in savanna-like habitats where some

of them even have synanthropic tendencies: P. urbicolum , true to its name, has often been found in

Dar-es-Salaam, and P. frundbergi has been collected in Mombasa and Tanga in Kenya (Enghoff et al.

2016). Some Prionopetalum species are even known as pests on various crops, including P. etiennei on

potato, African eggplant and okra in Senegal (Demange 1982; Etienne et al. 1992) and Prionopetalum

sp. (possibly P. xerophilum) on sweet potato in Uganda (Ebregt et al. 2005).

It is worth noting that P. kraepelini , the most Towlandistf Udzungwa odontopygid known so far, is

the only species among the Udzungwa odontopygids which (according to present knowledge) is not

endemic to this mountain range.

Acknowledgements

Thanks are due to the Tanzanian authorities that granted permits for collecting in the Udzungwa

Mountains, as well as to my colleagues Thomas Pape and Nikolaj Scharff for providing most of the

specimens, to Judith Winston and her colleagues at VMNH for access to specimens in that museum,

to Matthias Glaubrecht and Thure Dalsgaard for access to specimens in ZMUH, to Nesrine Akkari for

great help during and after my visit to NHMW, including provision of original images, and to Hans Reip

for access to old literature.

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Correction to Enghoff (2016)

On p. 17 of the previous article in the “A mountain of millipedes” series (Enghoff 2016), the reference

to Dieudonne (2016) should instead be to Ntashavu (2016).

Manuscript received: 3 February 2016

Manuscript accepted: 4 March 2016

Published on:22 July 2016

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, Fondon, United

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

Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Feiden, the Netherlands.