http://dx.doi.org/10.5852/eit.2014.10Q

www. europeani ournaloftaxonomv. eu

2014 • Enghoff H.

This work is licensed under a Creative Commons Attribution 3.0 License.

Monograph

urn:lsid:zoobank.org:pub:B3E6C489-6D96-4AF5-A33D-EE8329A9321B

A mountain of millipedes I: An endemic species-group of the genus

Chaleponcus Attems, 1914, from the Udzungwa Mountains, Tanzania

(Diplopoda, Spirostreptida, Odontopygidae)

Henrik ENGHOFF

Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100

Kobenhavn 0, Denmark, email: henghoff@snm .ku.dk

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

Abstract. Twenty new species of the millipede genus Chaleponcus Attems, 1914, are described from

the Udzungwa Mountains: C. netus sp. nov., C. quasimodo sp. nov., C. malleolus sp. nov., C. scopus sp.

nov., C. nikolajscharffi sp. nov., C. mwanihanensis sp. nov., C. basiliscus sp. nov., C. krai sp. nov., C.

nectarinia sp. nov., C. circumvallatus sp. nov., C. ibis sp. nov., C. vandenspiegeli sp. nov., C. vilici sp.

nov., C. teres sp. nov., C. hamerae sp. nov., C. termini sp. nov., C. gracilior sp. nov., C. mwabvui sp. nov.,

C. howelli sp. nov. and C. tintin sp. nov. Together with C. dabagaensis Kraus, 1958, they constitute the

Chaleponcus dabagaensis-group, well characterized by apparently apomorphic gonopodal characters,

presumably monophyletic, and the first example of a major radiation within the Udzungwas. All species

are restricted to altitudes >1390 m, all but one were found in only one, rarely two forest reserves, and

the vast majority of specimens were collected in montane forest. Chaleponcus gracilior sp. nov. was

collected in four forest reserves, often in secondary habitats where other species were only exceptionally

found. Co-occurrence of multiple species, inter-specific differences in body size and unusual tarsal

setation of a few species tentatively suggest adaptive radiation.

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

Enghoff H. 2014. A mountain of millipedes I: An endemic species-group of the genus Chaleponcus Attems, 1914,

from the Udzungwa Mountains, Tanzania (Diplopoda, Spirostreptida, Odontopygidae). European Journal of

Taxonomy 100: 1-75. http://dx.doi.org/10.5852/eit.2014.100

Contents

Introduction .2

Material and methods .4

Results .8

Taxonomy .8

General description of the Chaleponcus dabagaensis group.10

Species in which the gonopod coxa has an obvious lateral process.17

Chaleponcus netus sp. nov.17

Chaleponcus dabagaensis Kraus, 1958 .20

Chaleponcus quasimodo sp. nov. .23

European Journal of Taxonomy 100: 1-75 (2014)

Chaleponcus malleolus sp. nov.25

Chaleponcus scopus sp. nov.27

Chaleponcus nikolajscharjfi sp. nov.30

Chaleponcus mwanihanensis sp. nov.32

Chaleponcus basiliscus sp. nov.35

Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss)

is long and projects ± mesad.37

Chaleponcus krai sp. nov.37

Chaleponcus nectarinia sp. nov.39

Chaleponcus circumvallatus sp. nov.41

Chaleponcus ibis sp. nov.44

Chaleponcus vandenspiegeli sp. nov.46

Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss)

is not long and projecting ± mesad.48

Chaleponcus vilici sp. nov.48

Chaleponcus teres sp. nov.51

Chaleponcus hamerae sp. nov.53

Chaleponcus termini sp. nov.56

Chaleponcus gracilior sp. nov.58

Chaleponcus mwabvui sp. nov.61

Chaleponcus howelli sp. nov.63

Chaleponcus tintin sp. nov.65

Key to species of the Chaleponcus dabagaensis group.67

Distribution and habitat .69

Two notes on general morphology.70

Discussion .73

Acknowledgements.73

References .73

Introduction

Odontopygidae is an endemic Afrotropical family of millipedes counting around 400 described species,

all but one described from the African mainland. The exception, Bandeirenica caboverdus (Pierrard,

1987) was described from the Cape Verde Islands, where it is a significant pest (Jolivet 1986, Nascimento

2012) but it has most likely been introduced from the African mainland. Odontopygidae are thus absent

from Madagascar, like the two other major endemic Afrotropical millipede families, Oxydesmidae

Cook, 1895 (Hoffman 1990) and Gomphodesmidae Cook, 1896 (Hoffman 2005a).

Thanks to Kraus (1960, 1966) it is relatively easy to get an overview of the Odontopygidae, but the

family still poses major taxonomical problems; see e.g., Hoffman (1991) and Frederiksen & Enghoff

(2012). Up till now, 57 species of Odontopygidae have been described or recorded from Tanzania

(Enghoff et al. in prep.), but unidentified material already present in collections suggests that this is

just a small fraction of the actual number of species. The material from the Udzungwa Mts studied by

me alone counts 35 species. No less than 21 of these can be referred to the genus Chaleponcus Attems,

1914. Twenty species have been described in Chaleponcus until now (Kraus 1960, 1966; Frederiksen

2013a), mainly from southern Africa north to Zimbabwe and southern Mozambique. One species, C.

dabagaensis Kraus, 1958, was described from Tanzania but was later (Kraus 1960) excluded from the

genus. It was not until very recently that Frederiksen (2013a) “reintroduced” Chaleponcus to Tanzania

by describing a new species from the Pare Mts in the northern part of the country.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Genus concepts in Odontopygidae are to a high degree less than satisfactory (Frederiksen 2013b).

Add to this the fact that the identity of the type species of the type genus, Odontopyge Brandt, 1841,

was relatively recently (Hoffman 1991) totally re-interpreted, resulting in a complete nomenclatural

remix and leaving a considerable number of species described in Odontopyge without a valid generic

affiliation. Fuckily, Chaleponcus was not affected by this.

The present contribution is a taxonomic treatment of the 21 species of Chaleponcus so far collected

in the Udzungwa Mountains in southern Tanzania. The Udzungwa Mountains are part of the Eastern

Arc Mountains, which in turn constitute one among 25 biodiversity hotspots regarded as crucial for

conservation of biodiversity on a global scale (Myers et al. 2000); see also Burgess et al. (2007).

The Udzungwa Mountains, situated at T2T S to 8°38’ S and 35°07’ E to 36°58’ E (Fig. 1) and covering

more than 16,000 km 2 , is by far the largest among the eastern Arc mountain blocks. More than 1300 km 2

forest remains, spanning altitudes from 300 to 2580 m asl. The Udzungwas harbour an extremely rich

endemic fauna including 17 vertebrate species (Burgess et al. 2007).

There are several forest reserves in the Udzungwa Mts. Fig. 1 shows the location of those forest reserves

(henceforward abbreviated FR) from which material for the present paper has been collected.

Several millipede species have been reported from the Udzungwa Mountains (Enghoff 2011a; Enghoff

et al. in prep.; Hoffman 1990, 2005a, 2005b; Hoffman & Howell 1985, 1996; Kraus 1958). Almost

MIKUMI

Kisinga/

Rugaro

IK1NGA,

New Dabaga/

Ulangambi

Udzungwa

Scarp

W Kilombero Ndunduiu

Scarp

Nyambanito

Ukami

Kiranzi /

Kitungulu

Mwanihana

O Kigogo

20 km

Wooded grassland

Woodland

Open forest

Closed forest

O Towns

S Major roads

9^ Rivers

Fig. 1. Map of the Udzungwa Mountains, showing the location of the forest reserves (green rings) where

the material studied here was collected, as well as names of individual mountains in West Kilombero

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

European Journal of Taxonomy 100 : 1-75 ( 2014 )

all of these are as far as known endemic to the Udzungwas. The only odontopygid recorded from the

Udzungwas so far is the abovementioned Chaleponcus dabagaensis.

Marshall et al. (2001a, 2001b) analysed the millipede fauna of West Kilombero Scarp FR and New

Dabaga/Ulangambi FR, respectively, based on very large collections made by Frontier Tanzania which

were sorted into unnamed morphospecies. Marshall et al. (2001b) recognized 38 morphospecies of

millipedes from West Kilombero Scarp FR and 13 from New Dabaga/Ulangambi FR, but this is almost

certainly an underestimate of the actual number of species present in the analysed material. For example,

I regard it likely that most of the six Chaleponcus species described here from West Kilombero Scarp

FR would have been lumped as just one morphospecies, and similarly for most of the 11 Chaleponcus

species described here from New Dabaga/Ulangambi FR.

Material and methods

The vast majority of material for this paper comes from the zoological collections of the Natural History

Museum of Denmark, University of Copenhagen (ZMUC). Much of the material was collected during

several field trips to the Udzungwa Mts by ZMUC staff, but a very substantial part of it was collected

by the NGO Frontier Tanzania ( http://www.trontiergap.com/About-Us/Background-Mission.aspx l

and was subsequently deposited in ZMUC. Smaller collections of odontopygids from the Udzungwas

were obtained from the Virginia Museum of Natural History (VMNH), where a very large collection

of Tanzanian millipedes was accumulated by the late Richard L. Hoffman, and from Museo di Storia

naturale - Zoologia ‘Ta Specola’, Firenze (MZF).

Marshall et al. (2001a) described how the material from West Kilombero FR was collected: “Ground¬

dwelling millipedes were primarily sampled by timed searching of 3 m G m quadrats. ... The leaf

litter and topsoil within quadrats was searched thoroughly by hand for a total of eight person hours

per quadrat at all trapsites. In addition, the general proximity of all trapsites was searched for two

person hours. During such searches, particular attention was paid to rotting logs, the underside of rocks

and other such microhabitats, which may have been missed by the quadrat samples.” The collecting

procedure in New Dabaga/Ulangambi FR was the same (Marshall et al. 2001b). A. Marshall (pers.

comm.) has further informed me that during these campaigns the collectors had to dig down to around

15cm to find millipedes after initially fruitless searches in the topsoil alone.

A total of 460 male specimens were examined. All samples are kept in 70% alcohol, and the colours of

almost all specimens are quite bleached.

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 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.

Descriptions

In the light of the very large material at hand, of the uniformity of odontopygid millipede non-gonopod

morphology, and of the virtual impossibility of identifying females, only adult males are considered,

and the descriptions of non-gonopod morphology are kept to a minimum. The following information is

included:

• numbers of body rings (excluding telson)

• midbody diameter

• colour pattern when discernible

• limbus structure

• shape of anal valves

• setation of walking leg tarsi

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

The subheading “Distribution and habitat” summarises the label information. Altitudinal ranges are

given as the ‘maximal’ range although the actual range may be narrower. For example, the studied

samples of C. nectarinia sp. nov. were collected at 1955 m asl, 1945 m asl, 1940 m asl and 1930-1950 m

asl. The altitudinal range for this species is given as 1930-1955 m asl, although the specimens labelled

“1930-1955 m asl” may have been collected within the range specified by the exact altitudes given

for the other samples. It should be noted that the ‘maximal’ range given neglects a possible disjunct

altitudinal distribution. For example, for C. netus sp. nov. the ‘maximal’ altitudinal range 1390-1955 m

asl is given, but the samples are grouped in two ranges, viz., 1390-1410 m asl and 1800-1963 m asl.

Under the subheading “Coexisting species”, species found in the same samples as the species under

consideration, as well as additional species found in the same forest reserve, are listed; see also Table 1.

This information should be read keeping in mind that sampling was not at all standardized, and that a

sample may contain from one to 61 Chaleponcus males.

Gonopod terminology

The terminology introduced by Frederiksen (2013a,b; Frederiksen & Enghoff 2012) is used. This

terminology largely refrains from attempting to homologize parts of the gonopod telopodite with

individual podomeres (femur, tibia, tarsus ...) as was done by previous authors working on the group,

notably Kraus (1960, 1966).

Positional/directional terms:

Basal refers to a position on the side nearest to the gonopod sternum. Basad means “directed towards

the sternum”. Morphologically this refers to “dorsal” and “dorsad”, but since gonopods are traditionally

depicted upside down, basal and basad are preferred.

Apical refers to a position farthest away from the gonopod sternum. Apicad means “directed away

from the sternum”. Morphologically this refers to “ventral” and “ventrad”, but since gonopods are

traditionally depicted upside down, apical and apicad are preferred.

Distal refers to a position at or near the tip of the gonopod coxa, solenomere or telomere. Distad refers

to the corresponding directions. The antitheses are proximal and proximad .

Anatomical terms

The gonopod coxa consists of a longitudinally folded lamella with the free margins projecting mesad.

The anterior lamella is called proplica , the posterior lamella, metaplica. In Chaleponcus , proplica and

metaplica come together apically and form a hoodlike cucullus. See Fig. 7.

The gonopod telopodite consists of three main parts: basomere, solenomere and telomere. The basomere

starts as a straight rod, located in the cavity formed by the coxal pro- and metaplica. It then bends mesad;

the bend is termed arculus and is effectuated by a VA turn spiralling of the basomere whereby the spiral

loops are displaced mesad (somewhat like a high-spired snail shell). The part of the basomere involved

in the VA turn is called the torsotope. The actual right-angle bend takes place after the torsotope when

the basomere is “on its way down” (basad) but then bends mesad at right angles and becomes narrower

(posttorsal narrowing sensu Frederiksen 2013b). In the terminology of Kraus (1966), the part of the

basomere before the ‘actual’ bend is the prefemur, the posttorsal narrowing is the femur. After the

posttorsal narrowing the telopodite is divided into solenomere and basomere. The ‘postfemoral’ part

sensu Kraus (1966) is not well-defined. The start of the solenomere can be difficult to define, but the fine

suture running along the solenomere (visible, e.g., on Fig 28B) helps to identify it. See Figs 8 and 10.

European Journal of Taxonomy 100 : 1-75 ( 2014 )

Table 1. Coexistence of Chaleponcus species in the UdzungwaMts. For each species, coexisting species

at two levels are listed: species found in the same samples and additional species found in the same forest

reserve. Abbreviations: FR = forest reserve, KIG = Kigogo, KK = Kiranzi/Kitunguli, KR = Kisinga/

Rugaro, MWA = Mwanihana, NDU = New Dabaga/Ulangambi, US = Udzungwa Scarp, WKS = West

Kilombero Scarp.

species

in same samples

further species in same FR

NDU, WKS

netus

NDU: dabagaensis, gracilior,

malleolus, mwabvui, nectarinia,

termini, vandenspiegeli, vilici

WKS: circumvallatus

NDU: lcrai, teres

WKS: basiliscus, gracilior, ibis,

tintin

KK, NDU

dabagaensis

KK: gracilior

NDU: gracilior, mwabvui, netus,

termini

KK: -

NDU: krai, malleolus, nectarinia,

teres, vandenspiegeli, vilici

KIG

quasimodo

howelli, mwabvui, scopus

NDU

malleolus

gracilior, nectarinia, netus, vilici

dabagaensis, krai, mwabvui, teres,

termini, vandenspiegeli

KIG

scopus

howelli, mwabvui, quasimodo

nikolajscharffi

hamerae

circumvallatus

MWA

mwanihanensis

WKS

basiliscus

ibis

circumvallatus, gracilior, netus,

tintin

KR, NDU

krai

KR: -

NDU: gracilior

KR: gracilior

NDU: -

NDU

nectarinia

malleolus, netus, teres, termini,

vilici

dabagaensis, gracilior, krai,

mwabvui, vandenspiegeli

US, WKS

circumvallatus

US: -

WKS: netus

US: hamerae, nikolajscharjfi

WKS: basiliscus, gracilior, ibis,

tintin

WKS

ibis

basiliscus

circumvallatus, gracilior, netus,

tintin

NDU

vandenspiegeli

gracilior, netus, termini

dabagaensis, krai, malleolus,

mwabvui, nectarinia, teres, vilici

NDU

vilici

gracilior, malleolus, nectarinia,

netus, teres, termini

dabagaensis, krai, mwabvui,

vandenspiegeli

NDU

teres

termini, vilici

dabagaensis, gracilior, krai,

malleolus, mwabvui, nectarinia,

netus, vandenspiegeli

hamerae

nikolajscharffi

circumvallatus

NDU

termini

gracilior, mwabvui, nectarinia,

netus, teres, vilici

dabagaensis, krai, malleolus,

vandenspiegeli

KK, KR,

NDU, WKS

gracilior

KK: dabagaensis

KR: -

NDU: dabagaensis, krai,

malleolus, mwabvui, netus,

termini, vilici

WKS: -

KK: -

KR: krai

NDU: nectarinia, teres,

vandenspiegeli

WKS: basiliscus, circumvallatus,

ibis, netus, tintin

KIG, NDU

mwabvui

KIG: howelli, quasimodo, scopus

NDU: dabagaensis, gracilior,

netus, termini

KIG: -

NDU: krai, malleolus, nectarinia,

teres, vandenspiegeli, vilici

KIG

howelfi

mwabvui, quasimodo, scopus

WKS

tintin

basiliscus, circumvallatus,

gracilior, ibis, netus

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Abbreviations for morphological terms used in descriptions and on illustrations

Accessory structures may be indicated with ' or with numbers.

anterior telomeral lamella

alp

process on al

als

spine on al

bal

basal telomeral lamella

cucullus

distal telomeral process

intermediate telomeral lamella

metaplical incision

protruding lateral margin of coxa

lateral coxal process

anteriad metaplical flange

mesal metaplical process

mfp

metaplical flange process

metaplical shelf

msl

metaplical shelf-forming lobe

mss

metaplical shelf-spine

proximal side branch of slm

posterior telomeral lamella

plb

basal branch of pi

pip

process on pi

pis

spine on pi

prl

proplical lobe

proximal solenomeral spine

pxl

proximal telomeral lobe

pxs

spine on proximal telomeral lobe

slm

solenomere

spl

spine-like telomeral branch

main telomeral stem

sts

.small spine on st

thumblike process on main telomeral stem

torsotope process

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

asl

MZF

VMNH

ZMUC

above sea level

Forest Reserve

Zoological Museum, Firenze

Virginia Museum of Natural History

Natural History Museum of Denmark (Zoological Museum)

European Journal of Taxonomy 100 : 1-75 ( 2014 )

Results

Taxonomy

Class Diplopoda Blainville-Gervais, 1844

Order Spirostreptida Brandt, 1833

Family Odontopygidae Attems, 1909

Subfamily Archepyginae Manfredi, 1939

The taxon Archepyginae corresponds to Odontopyginae sensu Kraus (1960, 1966), see Hoffman (1991),

and includes those odontopygid genera in which the gonopod telopodite has a flexible, mostly spiralled

zone (torsotope sensu Hoffman 2008) followed by a delimited, usually narrower zone (“Femur” sensu

Kraus 1960, 1968).

Tribe Prionopetalini Hoffman, 1991

This taxon was introduced as a replacement name for Odontopygini sensu Hoffman (1980) and includes

those archepygine genera in which the solenomere is long and thin, as opposed to the genera placed by

Hoffman (1980, 1991) in the tribe Ctenoiulini.

Genus Chaleponcus Attems, 1914

This genus was established by Attems (1914) for three new species, of which C. limbatus Attems, 1914,

was subsequently designated as type species by Kraus (1960).

Chaleponcus was treated in detail by Kraus (1960, 1966) and was recently addressed by Frederiksen

(2013a). Vohland & Hamer (2013) redescribed two Chaleponcus species from Namibia. From the works

of these authors the following diagnosis of Chaleponcus can be extracted, based entirely on gonopod

structure, using the terminology suggested by Frederiksen (2013a,b; Frederiksen & Enghoff 2012) and

emended in light of the species described here:

Prionopetalini in which

• the proplica and metaplica of the gonopod coxa come together apically and form a hoodlike structure,

here termed cucullus

• the solenomere is very long, whiplike, at least twice as long as the telomere if stretched out, not

spiralled terminally, and without accessory branches or outgrowths (except for sometimes at the

very base, corresponding to the “Tibialdorn” of Kraus; see species descriptions)

• the telomere is proximally folded like a tube or a trough and distally separates into two or three

diverging lamellae

Kraus (1960, 1966) included 19 species in Chaleponcus , distributed in southern African as far north

as Zimbabwe and southern Mozambique, and Frederiksen (2013a) added a 20 th species from northern

Tanzania. Kraus (1960, 1966) did not include Chaleponcus dabagaensis Kraus, 1958, described from

the Udzungwa Mts (not W Usambaras as stated by Frederiksen 2013a); this species Kraus (1960)

excluded from the genus and tentatively assigned to another predominantly S African genus, Spinotarsus

Attems, 1909. In fact, he described the species as Chaleponcus (,Storthoporus ) dabagaensis , and as he

(1960) listed Storthoporus Attems, 1928, as a synonym of Spinotarsus, the transfer seemed logical.

However, C. dabagaensis lacks most characteristics of Spinotarsus , including a spiny proximal lamella

on the telomere and the strongly sclerotized lamella (“Bogenlamelle”) on the posterior surface of the

telomere. C. dabagaensis does have a trait seen in many Spinotarsus species, viz., the longitudinally

fluted solenomere; this is a character not seen in other previously described Chaleponcus species but

found in most of the species described here. C. dabagaensis also has the spine-bearing shelf, see below,

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

and without doubt belongs together with the other Udzungwan species treated in the present paper. I here

reallocate it back to Chaleponcus.

The Udzungwa species described here and referred to Chaleponcus differ from their congeners in one

conspicuous character, the spine-bearing ‘shelf’ on the gonopodal coxal metaplica. Many odontopygids

have spines in various positions on the gonopod coxa, including the subdistal position on the metaplica

where the spine-bearing shelf is situated in the Udzungwan species. This is, e.g., true for Chaleponcus

parensis Frederiksen, 2013, Prionopetalum bifidum VandenSpiegel & Pierrard, 2009, Patinatius attemsi

Kraus, 1960, P. bidentatus Kraus, 1960, Spinotarsus viridis Kraus, 1966, Allantogonus spp. (Kraus

1960, Attems 1935 ), Kompsoprium spp. (Kraus 1960, Attems 1935) and Odontopygista natal ica Kraus,

1960. In all these species except one, however, the spine is not associated with a shelflike structure and

are most likely not homologous with the ‘shelf-borne’ spine in the Udzungwan species. The exception

is Spinotarsus viridis , where the spine does seem to originate from a horizontal shelf (Kraus 1966: fig.

350); in other characters, however, S. viridis is a typical Spinotarsus.

Based on the probably autapomorphic spine-bearing shelf and the longitudinally fluted solenomere,

a species group can be defined for the Udzungwan species of Chaleponcus. Following tradition, the

group is named after the first described species in the group and hence wifi be named the Chaleponcus

dabagaensis group.

3,5

-*—■

2,5

1,5

:: malleolus

A tintin

♦ gracilior

O netus

X mwanihanensis

• scopus

□ quasimodo

A nikolajscharffi

* dabagaensis

♦ circumvallatus

A krai

• basiliscus

■ ibis

“I

Number of podous rings

Fig. 2. Body size of males of 13 species of the Chaleponcus dabagaensis group: Number of podous

rings and vertical midbody diameter in large, small and Tong, thin’ species (cf. text), as well as medium

sized species in which the gonopod coxa has an obvious lateral process.

European Journal of Taxonomy 100: 1-75 (2014)

General description of the Chaleponcus dabagaensis group

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

includes a selection (cf above).

Non-gonopodal characters

Body length, ca. 2-5 cm. Midbody vertical diameter 1.6-3.5 mm. 38-54 podous rings, no apodous

rings in front of telson. The 21 species may conveniently be divided into four size groups (cf Figs 2-3):

• Large species, diameter 2.8-3.5 mm, 48-54 podous rings: C. basiliscus sp. nov., C. circumvallatus

sp. nov., C. ibis sp. nov., C. krai sp. nov.

• Medium-sized species, diameter 1.6-2.4 mm, 41M9 podous rings: C. dabagaensis, C. hamerae sp.

nov., C. howelli sp. nov., C. mwabvui sp. nov., C. mwanihanensis sp. nov., C. nectarinia sp. nov.,

C. netus sp. nov., C. nikolajscharffi sp. nov., C. quasimodo sp. nov., C. scopus sp. nov., C. teres sp.

nov., C. termini sp. nov., C. vandenspiegeli sp. nov., C. vilici sp. nov.

• Small species, diameter 1.6-1.8 mm, 38—41 podous rings: C. malleolus sp. nov., C. tintin sp. nov.

• ‘Long, thin species’, diameter 1.5-1.8 mm, 45M9 podous rings: C. gracilior sp. nov.

Colour. Bleached in most studied specimens, but a pale middorsal longitudinal band is often discernible.

Limbus (examined on the dorsal side of midbody rings). Lobulate to serrate. Lobes often triangular and

pointed, but sometimes rounded or spatulate, sometimes very shallow, rarely virtually absent, often

longitudinally striolate, sometimes apically denticulate. See Fig. 4.

Claws of walking legs. With a long (> claw proper) accessory claw. Tip of tarsus usually with a moderate

number of strong setae (“normal setation”), but in two species, C. circumvallatus sp. nov. and C. ibis sp.

nov., with a dense whorl of stout setae surrounding the claw like a palisade. See Fig. 5.

Telson. Preanal ring with wrinkled/coriaceous sculpture dorsally. Anal valves with wrinkled/coriaceous

sculpture, except along caudal and ventral margins; in almost all species each with a dorsal denticle of

variable size, and often also with a smaller ventral denticle; denticles when present well set off from rest

•*—>

l + 1

+ j

X ♦ ♦

j x nxo g* o o j

A n v a

O termini

• mwabvui

X howelli

□ teres

▲ vandenspiegeli

O vilici

♦ hamerae

+ nectarinia

U 9 U X O i

o © • • □

o • j

° !

\o /

-Z

35 37 39 41 43 45 47 49 51 53 55

Number of podous rings

Fig. 3. Body size of males of 8 species of the Chaleponcus dabagaensis group: Number of podous rings

and vertical midbody diameter in medium-sized species in which the gonopod coxa has no obvious

lateral process.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

of valve rather than just being “sharp corners” (compare Fig. 6A-C, with fig. 3 in Frederiksen 2013b).

Each valve with three setae. Free margin (‘lip’) of anal valves raised in most species and provided

with three small extensions (‘ravelins’) on which the setae are borne. These setiferous tubercles not

protruding in lateral view. See Fig. 6.

Body ring 7. Body ring 7 of males modified, with posterior shallow concavities for accommodation

of gonopods. The detailed shape of the concavities varies among species, in parallel with variations of

gonopod shape, but this has not been studied in detail here.

Gonopods

Coxa (Fig. 7). In anterior or posterior view 3M x as long as broad. Free margin of proplica straight,

shallowly concave or shallowly sinuous from base to proplical lobe (prl) at ca. % of the length of

the proplica. Metaplica basally with an anteriad flange {inf) usually ending in a blunt or triangular

process ( mfp). Free margin of metaplica variable, but at ca. % of its length, the metaplica carries a

metaplical shelf (ms) on which a metaplical shelf-spine (mss) is inserted. Distal to prl and ms , proplica

and metaplica join to form an apical ‘hood’, the cucullus ( cu). The cucullus may take many different

shapes and together with the metaplical shelf-spine defines the general ‘profile’ of the gonopod coxa

which is often species-specific (Fig. 9).

Telopodite (Figs 8, 10). Basomere including torsotope without spines, arculus 90°. First full (360°) turn

of torsotope very tight, leaving no space inside the spiral, but last half (180°) turn, a more open spiral,

leaving room for a triangular to semicircular torsotope process (tp) on the basal side of the spiral (tp was

described for C. dabagaensis by Kraus (1958) as a blunt peg in the position corresponding to that of a

femoral spine: “Kein Femoraldorn, auf der Vorderseite jedoch an dessen Stelle ein stumpfer Zapfen”).

The torsion of the torsotope begins with an anteriad bend. Posttorsal narrowing unremarkable, not very

pronounced.

Solenomere (slm). Solenomore originating on apical side of telopodite, just after posttorsal narrowing,

whiplike, taeniate, irregularly curving in specimens prepared for SEM, longitudinally fluted (sometimes

indistinctly) and sometimes also with tiny retrorse spines in distal part. A slender proximal spine (ps)

often originating from base of solenomere, sometimes accompanied by a small ± lamellar outgrowth

and/or a second small spine.

Telomere. Telomore highly variable, originating on basal side of telopodite, just after posttorsal

narrowing, starting with a proximal lobe (pxl) lying closely against posterior base of slm. Telomere

continuing distally to pxl in simple, large main stem (st), anterior margin of st forming shallow lobe

(.st ’) lying closely against proximal part of slm. Distal part of telomere very complicated, consisting of

several lamellar parts; two or three lamellae can be distinguished, but these may each have accessory

lamellae, carry variously shaped spines and have more or less serrated margins. I have been unable

to homologize the various lamellae etc. of the telomere between species; thus, what is called anterior

lamella in one species may be homologous with what it called posterior lamella in another. For example,

the very characteristic anterior lamella in C. krai sp. nov. is remarkably similar to the posterior lamella

in C. nectarinia sp. nov., but whether these two structures are homologous, or whether different parts of

the telomere have evolved convergently to serve similar needs, remains unclear.

Included species (alphabetically)

Chaleponcus basiliscus sp. nov., C. circumvallatus sp. nov., C. dabagaensis Kraus, 1958, C. gracilior

sp. nov., C. hamerae sp. nov., C. howelli sp. nov., C. ibis sp. nov., C. krai sp. nov., C. malleolus sp.

nov., C. mwabvui sp. nov., C. mwanihanensis sp. nov., C. nectarinia sp. nov., C. netus sp. nov., C.

European Journal of Taxonomy 100: 1-75 (2014)

nikolajscharjfi sp. nov., C. quasimodo sp. nov., C. scopus sp. nov., C. teres sp. nov., C. termini sp. nov.,

C. tintin sp. nov., C. vandenspiegeli sp. nov., C. vilici sp. nov.

Fig. 4. Limbus types in Chaleponcus species, SEM micrographs, external view, except I: internal view.

— A. Lobes pointed, more than twice as long as broad (C. scopus sp. nov.). B. Lobes pointed, slightly

less than twice as long as broad (C. howelli sp. nov.). C. Lobes pointed, almost equilateral triangles (C.

netus sp. nov.), D. Lobes rounded, slightly longer than broad (C. dabagaensis Kraus, 1958). E. Lobes

broadly rounded, slightly shorter than broad (C. hamerae sp. nov.). F. Lobes spatulate (C. gracilior sp.

nov.). G. Lobes rounded, broader than long, spinose, ridged (C. krai sp. nov.). H. Lobes ca. 3 times as

broad as high, rounded, spinose (C. teres sp. nov.). I. Lobes rounded, much broader than high, smooth (C.

malleolus sp. nov.). J. No lobes (C. circumvallatus sp. nov.). Scales: 0.01 mm (A-F, J), 0.001 mm (G-I).

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 5. Chaleponcus spp., tip of tarsus and claw of walking legs. — A. C. mwanihanensis sp. nov. as an

example of normal setation. B-C. C. circumvallatus sp. nov. with the claw surrounded by a ’palisade’

of setae. Scales 0.1 mm.

Fig. 6. Telson types in Chaleponcus species. —A. Dorsal and ventral anal valve spines well developed (C.

nectarinia sp. nov.). B. Spines very long (C. circumvallatus sp. nov.). C. Dorsal spine well developed, ventral

spine barely indicated (C. mwabvui sp.nov.). D.Nospines(C. teres sp.nov.). E. Setiferoustuberclesonravelins

(C. nectarinia sp. nov.). F. With setiferous tubercles, not on ravelins (C. mwabvui sp. nov.). Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

The species cannot easily be arranged in subgroups. For practical reasons the descriptions are organised

as follows:

• Species in which the gonopod coxa has an obvious lateral process (exceptionally, C. basiliscus

sp. nov., two processes): C. netus sp. nov., C. dabagaensis Kraus, 1958, C. quasimodo sp. nov., C.

malleolus sp. nov., C. scopus sp. nov., C. nikolajscharffi sp. nov., C. mwanihanensis sp. nov., C.

basiliscus sp. nov.

• Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss) is long

and projects ± mesad: C. krai sp. nov., C. nectarinia sp. nov., C. circumvallatus sp. nov., C. ibis sp.

nov., C. vandenspiegeli sp. nov.

• Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss) is not

long and projecting ± mesad (C. vilici sp. nov. has the lateral side of the coxa somewhat angled.

Fig. 7. Gonopod coxa terminology, left coxa of Chaleponcus termini sp. nov., mesal(-posterior) view, cu

= cucullus, mf= anteriad metaplical flange, mfp = metaplical flange process, ms = metaplical shelf, mss

= metaplical shelf-spine, prl = proplical lobe.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 8. Gonopod telopodite terminology. —A. Feft telopodite of Chaleponcus termini sp. nov., posterior

view. White oval = basomere, blue oval = torsotope, red oval = telomere, yellow = solenomere ( slm ),

green = proximal spine of solenomere {ps ), pxl = proximal lobe of telomere, st = main stem of telomere,

st ’ = shallow lobe of st. B-C. Torsotope (left gonopod) with torsotope process ( tp ). B. C. vilici sp. nov.,

mesal-anterior view. C. C. netus sp. nov., posterior view. D. Solenomere (C. dabagaensis) to show

longitudinal fluting. E. Tip of solenomere (C. netus sp. nov.) to show spinulation. Scales: 0.1 mm (A-

D), 0.01 mm (E).

European Journal of Taxonomy 100: 1-75 (2014)

which reminds of a small lateral process): C. vilici sp. nov., C. teres sp. nov., C. hamerae sp. nov.,

C. termini sp. nov., C. gracilior sp. nov., C. mwabvui sp. nov., C. howelli sp. nov., C. tintin sp. nov.

Fig. 9. Chaleponcus spp., outline of coxite of left gonopod, posterior view. (The ’transverse’ plane of

the gonopod is usually slightly at angles with the exact transverse plane of the millipede, so it may be

necessary to tilt the gonopod slightly in order to obtain the view shown on the drawings.) The stippled

lines indicate the place where the telopodital basomere emerges from the coxal cavity. Not to scale.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Species in which the gonopod coxa has an obvious lateral process

(exceptionally, C. basiliscus sp. nov., two processes)

Chaleponcus netus sp. nov.

urn:lsid:zoobank.org:act:244DB18B-A3BC-43C6-9B7F-BlC309C 14737

Figs 4, 8-11, 34

Diagnosis

Medium-sized. Differs from all other group members by the presence of a long, gently curved,

longitudinally fluted spine ( pxs ) on the proximal lobe of the telomere. The profile of the coxa (Fig. 9) is

also distinctive, as is the spine-like tip of the anterior lamella (al) of the telomere.

Etymology

The name is a Fatin adjective meaning ’’woven” or ’’ kni t” and refers to the intricately interwoven

gonopod telopodites.

Material studied (total: 66 33)

Holotype

TANZANIA: 3 , Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°00’

26.6” S, 35°56’ 06.1” E, montane forest, 1910 m asl, plot Kinyonga, casual, 24 Oct. 2000, Frontier

Tanzania (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, all collected by Frontier Tanzania,

all in ZMUC: New Dabaga/Ulangambi FR: 9 33 , 08°05’34.5” S, 35°55’ 31.5” E, montane, 1800-1900

m asl, plot 16, casual, 15-16 Nov. 2000; 3 33 , 08°00’ 26.6” S, 35°56’ 06.1” E, montane forest, 1910 m

asl, plot Kinyonga, casual, 24 Oct. 2000; 1 3 , 08°00’ 26.5” S, 35°56’ 06.1” E, scrub/thicket/bush, 1908

m asl, plot Kinyonga, 19 Oct. 2000. West Kilombero Scarp FR: 1 3 , 07°50’ 38.4” S, 36°22’ 17.6” E,

montane forest, 1390 m asl, plot Paradiso, 12 Nov. 2000; 6 33 , 07°50’ 38.4” S, 36°22’ 17.6” E, montane

forest, 1390-1410 m asl, plot Paradiso, 18 Nov. 2000; 2 33 , 08°04’ 05.6” S, 35°54’ 20.4” E, montane,

1930 m asl, plot 6, sample 1 of 3, 28 Oct. 2000; 2 33 , 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930

m asl, plot 6, sample 2 of 3, 27 Oct. 2000; 1 3 , 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930 m asl,

plot 6, sample 3 of 3, 27 Oct. 2000; 2 33 , 08°03’ 39.9” S, 35°54’ 41.9” E, montane, 1945 m asl, plot

24, 29 Oct. 2000; 16 33 , 08°03’ 34.9” S, 35°54’ 41.9” E, montane, 1955 m asl, plot 24, casual, 2 Nov.

2000; 2 33 , 08°03’ 39.9” S, 35°54’ 41.9” E, montane, 1963 m asl, plot 24, sample 1 of 3, 28 Oct. 2000;

3 33 , montane, 1800-1955 m asl, plot MS, plot A, 29 Oct. 2000; 17 33 , 07°50’ 38.4” S, 36°22’ 17.6”

E, montane forest, 1390-1410 m asl, plot Paradiso, 12-18 Nov. 2000.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°00’ 26.6”

S, 35°56’ 06.1” E, montane forest, 1910 m asl.

Description (male)

Diameter. 1.9-2.3 mm, 41—45 podous rings.

Colour. After 12 years in alcohol pale yellowish, with paramedian dark bands flanking a broad middorsal

light stripe, a narrow blackish lateral stripe at ozopore level.

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 10. Chaleponcus netus sp. nov., left gonopod, coloured. — A. Posterior view. B. Ventral view.

Orange = coxite, blue = torsotope, yellow = solenomere ( slm ), green = proximal spine of solenomere

( ps ), pink = telomere, pxs = spine on proximal telomeral lobe, st = main stem of telomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 11. Chaleponcus netus sp. nov., gonopods. — A. Gonopods, anterior view. B. Feft coxite, mesal

view. C. Base of proximal solenomere spine (ps ) with spinose socket. D. Right gonopod, anterior (and

slightly mesal) view. E. Feft gonopod telopodite, posterior view. F. Base of proximal solenomere spine

(ps) in a different individual, with denticle (ps2). al = anterior lamella of telomere, il = intermediate

lamella of telomere, mfp = metaplical flange process, ms = metaplical shelf, mss = metaplical shelf-

spine, pi = posterior lamella of telomere, ps = proximal spine of solenomere, ps2 = additional proximal

spine of solenomere, pxl = proximal lobe of telomere, pxs = spine on proximal lobe of telomere. Scales:

0.1 mm (A-B, D-E), 0.05 mm (C, F).

European Journal of Taxonomy 100: 1-75 (2014)

Anal valves. Each with a long, slightly curving dorsal spine and a small to well-developed, triangular

ventral one; marginal rim raised, setiferous tubercles on small ‘ravelins’.

Limbus (Fig. 4C). With almost equilateral triangular lobes with rounded, spinulose tips, separated by

parabolic sinuses; lobes striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9-11). About 4 x as long as wide. Lateral margin very shallowly convex from base

up to beyond level of prl and ms , then curving lateral and together with apical margin forming blunt,

disto-lateral process of cucullus; main part of cucullus subrectangular in anterior view. Metaplical flange

(mf) ending in blunt process; metaplical mesal margin straight up to level of arculus, then abruptly

turning mesad at ca. 130° angle (base of mss), then turning apicad at ca. right angles. Metaplical shelf

(ms) regularly rounded, projecting posteriad as well as mesad. Metaplical shelf-spine (mss) originating

from anterior-mesal end of ms, very long, first directed apicad, then mesad and apically slightly obliquely

distad, in situ crossing over with opposite mss.

Gonopod telopodite (Figs 8, 10-11). Solenomere with a very long, gently curved proximal spine (ps)

set in a socket with more or less spinulose rim; sometimes a tiny denticle (ps2) at base of ps. Proximal

lobe of telomere (pxl) with a long, gently curved, longitudinally fluted spine (pxs). Telomere distally

with three branches:

• an S-shaped anterior lamella (al), broad at base, then narrowing and ending in curved spine-like tip,

• a large, irregularly shaped intermediate lamella (il),

• a small, rounded posterior lamella (pi).

Distribution and habitat

Known from New Dabaga/Ulangambi FR and West Kilombero FR, Udzungwa Mts. Altitudinal range:

1390-1963 m asl. Habitat: montane forest; one sample from scrub/thicket/bush.

Coexisting species

In New Dabaga/Ulangambi FR C. dabagaensis, C. gracilior sp. nov., C. malleolus sp. nov., C. mwabvui

sp. nov., C. nectarinia sp. nov., C. termini sp. nov., C. vandenspiegeli sp. nov. and C. vilici sp. nov.

were found in the same samples as C. netus sp. nov. In addition, C. krai sp. nov. and C. teres sp. nov.

were found in New Dabaga/Ulangambi FR. In West Kilombero Scarp FR C. circumvallatus sp. nov.

was found in the same sample as C. netus sp. nov. In addition, C. basiliscus sp. nov., C. gracilior sp.

nov., C. ibis sp. nov., and C. tintin sp. nov. were found in West Kilombero Scarp FR.

Chaleponcus dabagaensis Kraus, 1958

Figs 4, 8-9, 12

Chaleponcus (Storthoporus) dabagaensis Kraus, 1958

Chaleponcus (Storthoporus) dabagaensis'. Kraus 1960: 192, under “Species incertae sedis”.

Diagnosis

Medium-sized. Differs from all other group members by the presence of a thumblike process (th) on the

main stem of the telomere; also by the profile of the gonopod coxa (Fig. 9), especially its ‘hunch-backed’

lateral side, shared only by C. quasimodo sp. nov.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, Kwabuni Estate at Dabaga, 2100 m asl.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Material studied (total: 9 33)

Type material

Holotype 3 (Uberseemuseum Bremen), paratypes 5 $ $ (Uberseemuseum Bremen and Senckenberg

Museum Frankfurt), all from type locality; not studied.

Other material

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, all in ZMUC: Kiranzi-Kitungulu

FR: 2 S3, 08°09’S, 35°05’E, forest, Jan. 1996, M. Andersen, P. Gravlund, A. Jakobsen leg; 5, S3,

08°07’S, 35°04’E, 1700 m asl, Mar.-Apr. 1996, F.F. Sorensen leg. New Dabaga/Ulangambi FR: 1 3,

08°06’47.4” S, 35°56’50.4” E, montane, 1855 m asl, plot MS - plot A, 11 Nov. 2000, Frontier Tanzania

UMPS leg.; 1 3, 08°06’47.4” S, 35°56’50.4” E, montane, 1800-1955 m asl, plot MS -plot A, 29 Oct.

2000, Frontier Tanzania leg.

Description (male)

[data in square brackets from Kraus (1958), translated]

Diameter. 2.4 [2.4] mm, 49 [49] podous rings.

Colour. [Horn yellow, dorsally with darker marbling and medially with broad, yellow, longitudinal

stripe; a blackish lateral stripe at ozopore level].

Anal valves. Each with a long, slightly curving dorsal spine and a small, triangular ventral one; marginal

rim raised; setiferous tubercles well developed, on small ‘ravelins’.

Fimbus (Fig. 4D). With tongue-shaped, apically rounded lobes separated by narrow sinuses, lobes almost

meeting at base, lobes striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 12). About 4 x as long as wide. Fateral margin straight up to level of ms ,

then curving mesad and then lateral, leaving a deep, rounded incision and giving the entire coxa a

‘hunchback’ profile; apical margin gently convex, laterally gently concave, forming long, slightly

curved lateral cucullar process. Metaplical flange ( mf) ending in approximately right-angled-rounded

corner; metaplical mesal margin basally straight, with rounded-triangular process (mp) just below level

of arculus. Metaplical shelf (ms) simply rounded, projecting posteriad as well as mesad, continuing

without transition in metaplical shelf-spine ( mms ); mms of medium length, curving first mesad, then

obliquely basad, tip reaching level of basomere.

Gonopod telopodite (Figs 8,12). Solenomere with a small, straight proximal spine (ps) with an associated

lobe (as in C. gracilior sp. nov., but simpler). Main stem of telomere (si) with a characteristic thumblike

process (th) distally. Telomere distally with two branches:

• a large, convoluted anterior lamella (al) with smooth edges, continuous with st on basal side of

telomere,

• a posterior-distal lamella (pi) forming direct distal continuation of st but separated from it by

rounded- triangular incision; margins of pi in part strongly serrated.

Distribution and habitat

Known from New Dabaga/Ulangambi FR and Kiranzi-Kitungulu FR, Udzungwa Mts. Altitudinal range:

1700-1955 m asl, 2100 m asl according to Kraus (1958). Habitat: montane forest. Recorded by Kraus

(1958) from “Halbregenwald” (= semi-rain forest).

European Journal of Taxonomy 100: 1-75 (2014)

Coexisting species

In Kiranzi-Kitungulu FR, C. gracilior sp. nov. was found in the same sample as C. dabagaensis. In New

Dabaga/Ulangambi FR C. gracilior sp. nov., C. mwabvui sp. nov., C. netus sp. nov. and C. termini sp.

Fig. 12. Chaleponcus dabagaensis Kraus, 1958, left gonopod. — A. Anterior view. B. Posterior view.

C. Mesal-ventral view. D. Mesal-dorsal view, al = anterior lamella of telomere, mf= metaplical flange,

mp = mesal metaplical process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior

lamella of telomere, ps = proximal spine of solenomere, st = main telomeral stem, th = thumblike

process of telomere main stem. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

nov. were found in the same samples as C. dabagaensis. In addition, C. krai sp. nov., C. malleolus sp.

nov., C. nectarinia sp. nov., C. teres sp. nov., C. vandenspiegeli sp. nov. and C. vilici sp. nov. were found

in New Dabaga/Ulangambi FR.

Notes

The original description by Kraus (1958) leaves no doubt that the specimens at hand (which include

topotypes) belong to C. dabagaensis. See under “Genus Chaleponcus ” for a review of the history of this

species.

Chaleponcus quasimodo sp. nov.

urn:lsid:zoobank.org:act:37A2F533-120D-45C5-8C49-B77A4C2DA088

Figs 9, 13

Diagnosis

Medium-sized. Differs from all other group members except C. dabagaensis by the ‘hunch-backed’

profile of the gonopod coxa (Fig. 9); differs from C. dabagaensis by the profile of the coxal tip and by

the absence of a thumblike process on the main stem of the telomere.

Etymology

The name is a noun, here in apposition, and refers to Quasimodo, the hunch-back of Notre-Dame, title

figure of the novel by Victor Hugo (1839).

Material studied (total: 9 SS)

Holotype

TANZANIA: <$, Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, 1700 m, 8-10 Oct. 1984,

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

Paratypes

All from TANZANIA, Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, all in ZMUC: 1 3,

1700 m, 8-10 Oct. 1984, pitfall traps in montane rain forest. N. Scharff leg.; 7 33, 1900 m, 7-15 Oct.

1984, pitfall traps in montane rain forest. N. Scharff leg.

Type locality

TANZANIA, Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, 1700 m, montane rain forest.

Description (male)

Diameter. 2.1-2.3 mm, 47—49 podous rings.

Colour. After 30 years in alcohol almost completely bleached, with traces of paramedian darker bands

flanking a broad middorsal light stripe, sometimes a thin dark line in middle of stripe.

Anal valves. Each with a long, slightly curving dorsal spine and a small, triangular ventral one; marginal

rim raised, setiferous tubercles on small ‘ravelins’.

Fimbus. With broadly rounded lobes, as long as broad or slightly longer, striate on outer surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 13). About 3 x as long as wide. Fateral margin straight up to level of ms ,

then curving mesad and then lateral, leaving a deep, rounded incision and giving the entire coxa a

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 13. Chaleponcus quasimodo sp. nov., left gonopod. A. Anterior view. B. Anterior-mesal view.

C. Ventral view. D. Posterior view, al = anterior lamella of telomere, mf= metaplical flange, mp =

mesal metaplical process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior lamella of

telomere, ps = proximal spine of solenomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

‘hunchback’ profile; apical margin slightly convex, irregularly undulate, forming ca. 70° angles with

lateral margin. Mesal margin of proplica with two mesad processes apically, one above, one below

metaplical shelf-spine (mss). Metaplical flange (inf) ending in approximately right-angled-rounded

corner, hardly projecting; metaplical mesal margin with rounded-triangular process (mp) just below

level of arculus, distally with triangular incision with coarsely nodose margin. Metaplical shelf (ms) of

moderate size, projecting slightly posteriad as well as mesad, continuing without transition in metaplical

shelf-spine (mss); mss of medium length, curving obliquely basad.

Gonopod telopodite (Fig. 13). Solenomere with medium long, a lm ost straight proximal spine (ps).

Telomere distally with two relatively small and simple branches:

• an anterior lamella (al),

• a posterior lamella (pi).

Distribution and habitat

Known only from Kigogo FR, Udzungwa Mts. Altitudinal range: 1700-1900 m. Habitat: montane rain

forest.

Coexisting species

All four species known from Kigogo FR, i.e., C. quasimodo sp. nov., C. scopus sp. nov., C. howelli sp.

nov. and C. mwabvui sp. nov., were found in the same samples.

Chaleponcus malleolus sp. nov.

um:lsid:zoobank.org:act:85393BDE-7999-4CD5-BFA2-2B21A022E2A8

Figs 4, 9, 14

Diagnosis

Small (smallest species in the C. dabagaensis group). Differs from all other group members, except C.

teres sp. nov., by the absence of a dorsal spine on each anal valve. Further differs from all other group

members by the profile of the gonopod coxal tip (Fig. 9), especially the strongly curved, hooklike lateral

process.

Etymology

The name is a Fatin noun, here in apposition, meaning Tittle hammer’, and refers to the gonopod coxa

profile.

Material studied (total: 14 SS)

Holotype

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°00’

26.6” S, 35°56’ 06.1” E, montane forest, 1910 m asl, plot Kinyonga, casual, 24 Oct. 2000, Frontier

Tanzania (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, all

collected by Frontier Tanzania, all in ZMUC: 1 08°00’ 26.6” S, 35°56’ 06.1” E, montane forest, 1910

m asl, plot Kinyonga, casual, 24 Oct. 2000; 9 $$, 08°00’ 26.6” S, 35°56’ 06.1” E, scrub/thiclcet/bush,

1904-1915 m asl, plot Kinyonga, 18-20 Oct. 2000; 3 SS, 08°03’ 39.9” S, 35°54’ 41.9’” E, montane,

1945 m asl, plot 24, 29 Oct. 2000.

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 14. Chaleponcus malleolus sp. nov., right gonopod. A. Anterior view. B. Mesal view, focus on

coxa. C. Posterior view. D. Anterior-mesal view. E. Mesal view, focus on telopodite. F. Telopodite,

ventral-posterior view, al = anterior lamella of telomere, cu = cucullus, mfp = metaplical flange process,

mp = mesal metaplical process, mss ; metaplical shelf-spine, pll, pl2, pl3 = lobes of posterior lamella of

telomere, st = main stem of telomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°00’ 26.6”

S, 35°56’ 06.1” E, montane forest, 1910 m asl.

Description (male)

Diameter. 1.7-1.8 mm, 38-39 podous rings.

Colour. After 13 years in alcohol pale yellowish, traces of dark dorsolateral bands and a pale middorsal

band.

Anal valves. Neither dorsal nor ventral spines; marginal rim not raised, setiferous tubercles small, not

on ‘ravelins’.

Fimbus (Fig. 41). With very low, smooth lobes, ca. 0.2 times as long as broad.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 14). Almost 4 x as long as wide; lateral margin very slightly convex up to level

of proplical lobe, then curving laterad and then basad, forming cucullar hook together with otherwise

almost straight apical margin; cucullus (at) short, with big lateral, basad, blunt-tipped strongly curved

hook. Metaplical flange ending in blunt, apicad process ( mfp ); metaplical mesal margin straight, but with

triangular process (mp) just below level of arculus. Metaplical shelf mainly projecting mesad, continuing

without transition in metaplical shelf-spine (mss), mss medium long, curving almost exclusively in

horizontal plane, first anteriad, then mesad.

Gonopod telopodite (Fig. 14). Solenomere without a spine or other secondary branches. Telomere

simple, distally with 2, closely adjoining branches:

• an oblong anterior lamella (al),

• a posterior lamella forming a direct distal continuation of main stem ( st ) and distally with three

short lobes: apical (pll) and intermediate (pl2 ) lobe, with irregularly jagged edges, basal lobe (pl3 )

smooth-edged, semicircular.

Distribution and habitat

Only known from New Dabaga/Ulangambi FR. Altitudinal range: 1904-1945 m asl. Habitat: montane

forest, scrub/thicket/bush.

Coexisting species

C. gracilior sp. nov., C. nectarinia sp. nov., C. netus sp. nov. and C. vilici sp. nov. were found in the

same samples as C. malleolus sp. nov. In addition, C. dabagaensis, C. krai sp. nov., C. mwabvui sp. nov.,

C. teres sp. nov., C. termini sp. nov. and C. vandenspiegeli sp. nov. and were found in New Dabaga/

Ulangambi FR.

Chaleponcus scopus sp. nov.

um:lsid:zoobank.org:act:A4A8ElFE-3CE6-4D04-AlEE-80A63ED9FB79

Figs 4, 9, 15

Diagnosis

Medium-sized. Differs from all other group members by the profile of the gonopod coxal tip (Fig. 9),

especially the shape of the lateral process. Very similar to C. nikolajscharffi sp. nov., sharing, i.a., a non-

transverse metaplical shelf and a groove for accommodating the solenomere formed by two telomere

European Journal of Taxonomy 100 : 1-75 ( 2014 )

lobes; differing from C. nikolajscharffi sp. nov. by being thinner and having a differently shaped lateral

coxal process.

Etymology

The name is a Latin noun in apposition and refers to the profile of the gonopod coxa, which somewhat

resembles the head profile of the hammerhead stork ( Scopus umbretta).

Material studied (total: 16 c?c?)

Holotype

TANZANIA: Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, 1700 m, 8-10 Oct. 1984,

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

Paratypes

All from TANZANIA, Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, all in ZMUC: 6

1700 m, 8-10 Oct. 1984, pitfall traps in montane rain forest, N. Scharff leg.; 9 1900 m, 7-15

Oct. 1984, pitfall traps in montane rain forest, N. Scharff leg.

Type locality

TANZANIA: Iringa Region, Mufindi District, Udzungwa Mts, Kigogo FR, 1700 m, 8-10 Oct. 1984,

montane rain forest.

Description (male)

Diameter. 1.9-2.0 mm, 47—49 podous rings.

Colour. After 30 years in alcohol with traces of paramedian darker bands flanking a broad middorsal

light stripe; dark bands divided into separate dots on each diplosegment in anterior part of body.

Anal valves. Each with a long dorsal spine and a well-developed, but small, triangular ventral one;

marginal rim raised, setiferous tubercles well developed, on ‘ravelins’.

Limbus (Fig. 4A). With very long, slender triangular lobes; lobes ca. 2A x as long as broad, almost

meeting at base, striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 15). About 3 ’A x as long as wide. Lateral margin basally slightly convex, curving

strongly laterad ca. at level of prl , forming semicircular incision, and continuing in slender, curved,

pointed lateral process (Ip); apical margin sloping apico-mesad from process, irregularly undulate

and with blunt-angled projection ca. 2 A from tip of Ip; cucullus (cu) mesally projecting as rounded-

subrectangular lobe with irregularly undulate margin. Metaplical flange (mf) with very low rounded

anteriad protrusion, no process; metaplical mesal margin basally straight, then overlaid by lamelloid

process (mp) covering arculus. Metaplical shelf not transverse as in most other species: posterior surface

of metaplica with large lobe ( msl ) roofing arculus; lobe concave on posterior face, folded anteriad and on

anterior face giving rise to metaplical shelf-spine (mss), mms of medium length, curving first obliquely

anteriad, then mesad, then obliquely basad, not reaching level of torsus.

Gonopod telopodite (Fig. 15). Solenomere with a long, gently sinuous, mesad, proximal spine (ps)

arising from a wrinkled area. Telomere distally with two branches:

• a very large anterior lamella ( al ), increasing in diameter towards tip, with a broadly rounded smooth-

edged tip and a small, thumblike process (al’); solenomere resting between al’ and main body of al..

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

• a small, rounded posterior lamella (pi) with irregular edges.

Distribution and habitat

Known only from Kigogo FR, Udzungwa Mts. Altitudinal range: 1700-1900 m asl. Habitat: montane

rain forest.

Fig. 15. Chaleponcus scopus sp. nov., right gonopod. A. Posterior view. B. Ventral-posterior view.

C. Mesal-posterior view. D. Anterior view, al = anterior lamella of telomere, aV = thumblike process of

al, cu = cucullus, Ip = lateral coxal process, mf= metaplical flange, mp = mesal metaplical process, msl

= metaplical shelf-forming lobe, mss = metaplical shelf-spine, pi = posterior lamella of telomere, ps =

proximal solenomeral spine, slm = solenomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Coexisting species

All four species known from Kigogo FR, i.e., C. quasimodo sp. nov., C. scopus sp. nov., C. howelli sp.

nov. and C. mwabvui sp. nov., were found in the same samples.

Note

Very similar to and probably closely related to C. nikolajscharffi sp. nov. from Udzungwa Scarp FR.

Chaleponcus nikolajscharffi sp. nov.

urn:lsid:zoobank.org:act:FA4BA44E-0547-4E9C-A4A4-781E2298A650

Figs 9, 16, 34

Diagnosis

Medium-sized. Differs from all other group members by the profile of the gonopod coxal tip (Fig.

9). Very similar to C. scopus sp. nov., sharing, i.a., a non-transverse metaplical shelf and a groove for

accommodating the solenomere formed by two telomere lobes; differing from C. scopus sp. nov. by

apparently having the solenomere-conducting groove formed by the anterior and the posterior telomeral

lamellae, instead of by the anterior lamella and a process from the latter, as well as by having a differently

shaped lateral coxal process.

Etymology

The name honours my colleague, arachnologist Nikolaj Scharff (ZMUC), collector of this and numerous

other Tanzanian millipedes.

Material studied (total: 11 S3)

Holotype

TANZANIA: 3, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village, 1500 m,

2-13 Nov. 1984, pitfall traps, montane rain forest. N. Scharff leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR: 1 S, above Chita village,

1450-1650 m, 4-9 Nov. 1984, pitfall traps, montane rain forest, N. Scharff leg. (ZMUC); 2 S3, above

Chita village, 1600-1650 m, 8-13 Nov. 1984, pitfall traps, montane rain forest, N. Scharff leg. (ZMUC);

2 33, above Chita village, 1400 m, 4-5 Nov. 1984, pitfall traps, montane rain forest, N. Scharff leg.

(ZMUC); 1 3 , above Chita village, 1500 m, 2-13 Nov. 1984, pitfall traps, montane rain forest, N.

Scharff leg. (ZMUC); 2 33, 11 km SE of Masisiwe village, Kihanga Stream, 1800 m, 08°22’ 05.7” S,

35°58’ 41.6” E, 17-27 May 1997, ZMUC & SI Exp. leg. (ZMUC); 2 33, nr. Masisiwe, 08°20’ 32” S,

35°58’03” E, 12-15 Jul. 2004, A. Sforzi & L. Bartolozzi leg. (MZF, n° Mag. 2695).

Type locality

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

montane rain forest.

Description (male)

Diameter. 2.2-2.4 mm, 48—49 podous rings.

Colour. After 10 years in alcohol with a broad light middorsal stripe formed by an hourglass-shaped

spot on each body ring; stripe flanked by brownish paramedian bands; older specimens unicolourous

light brown/pinkish.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Anal valves. Each with a long, slightly curving spine and a well-developed, triangular ventral one;

marginal rim raised, setiferous tubercles well developed, on ‘ravelins’.

Fig. 16. Chaleponcus nikolajscharffi sp. nov., right gonopod. A. Mesal view. B. Anterior view. C. Ventral

view. D. Posterior view, al = anterior lamella of telomere, cu = cucullus, Ip = lateral coxal process, mf=

metaplical flange, mp = mesal metaplical process, msl = metaplical shelf-forming lobe, mss = metaplical

shelf-spine,/?/ = posterior lamella of telomere, = proximal spine of solenomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Limbus. With long, slender triangular lobes; lobes ca. 2Vi x as long as broad, almost meeting at base,

striate on external surface.

Tarsal Setation. Normal.

Gonopod coxa (Figs 9, 16). About 4 x as long as wide. Lateral margin basally slightly convex, curving

strongly laterad ca. at level of prl , forming semicircular incision, and meeting apical margin in

irregularly knobbed laterad process (Ip); apical margin sloping apico-mesad from process, irregularly

undulate; cucullus (cu) transverse, mesally projecting as rounded lobe with irregularly undulate margin.

Metaplical flange (mf) ending in low, rounded process; metaplical mesal margin basally straight, further

distally overlaid by lamelloid process (mp) covering arculus. Metaplical shelf not transverse as in most

other species: posterior surface of metaplica with large lobe (msl) roofing arculus; lobe concave on

posterior face, folded anteriad and on anterior face giving rise to metaplical shelf-spine (mss), mms of

medium length, curving first anteriad, then mesad, then basad, reaching level of torsus.

Gonopod telopodite (Fig. 16). Solenomere with a long, straight, mesad proximal spine (ps) originating

from a wrinkled area. Telomere distally with two branches, al and pi, both curving anteriad, both broad,

in part with irregularly serrate edges, lying closely together and accommodating solenomere.

Distribution and habitat

Known only from Udzungwa Scarp FR. Altitudinal rang: 1400-1800 m asl. Habitat: montane rain forest.

Coexisting species

C. hamerae sp. nov. was found in the same samples as C. nikolajscharffi sp. nov. In addition, C.

circumvallatus sp. nov. was found in Udzungwa Scarp FR.

Notes

Very similar and probably closely related to C. scopus sp. nov. from Kigogo FR.

Very peculiar, mushroom-like inter-cytoscutal structures were observed with the scanning electron

microscope in this species; see “Two notes on general morphology” and Fig. 34.

Chaleponcus mwanihanensis sp. nov.

urn: lsid: zoobank, org: act: 84D1DB74-24F 4-44 A7-BF 3 8 -DD31AOFAAE15

Figs 5, 9, 17

Diagnosis

Medium-sized. Without a proximal spine on the solenomere,

sp. nov. and C. hamerae sp. nov. Differing from these species

group) by the profile of the gonopod coxa (Fig. 9).

Etymology

The name is a Latin adjective referring to the type locality.

Material studied (total: 17 SS)

Holotype

TANZANIA: $, Morogoro Region, Kilombero District, Udzungwa Mts, Mwanihana FR, 1800-1850 m,

pitfall traps, montane rain forest 25-29 Sep. 1984, N. Scharff leg. (ZMUC).

a character shared only by C. malleolus

(and indeed from all other species in the

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 17. Chaleponcus mwanihanensis sp. nov., gonopods. A. Feft gonopods, mesal-ventral view. B. Feft

gonopod, posterior view. C. Right gonopod, ventral view. D. Right gonopod, anterior view, al = anterior

lamella of telomere, cu = cucullus, Ip = lateral coxal process, mf= metaplical flange, ms = metaplical

shelf, mss = metaplical shelf-spine, pi = posterior lamella of telomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Paratypes

All from TANZANIA, Morogoro Region, Kilombero District, Udzungwa Mts, Mwanihana FR, all in

ZMUC: 3 above Sanje, 1800 m, pitfall trap, 18 Aug. 1982, M. Stoltze & N. Scharffleg.; 1 $, above

Sanje, 1800 m, litter, 18 Aug. 1982, M. Stoltze & N. Scharffleg.; 9 <$<$, 1800-1850 m, pitfall traps,

montane rain forest, 25-29 Sep. 1984, N. Scharffleg.; 3 1800-1850 m, litter in montane rain forest,

28-29 Sep. 1984, N. Scharffleg.

Type locality

TANZANIA: Morogoro Region, Kilombero District, Udzungwa Mts, Mwanihana FR, 1800-1850 m,

montane rain forest.

Description (male)

Diameter. 2.2-2.4 mm, 44^16 podous rings.

Colour. After 30 years in alcohol bleached but with traces of a broad, light dorsal stripe, flanked by dark

dorsolateral bands, and thin dark lines at ozopore level.

Anal valves. Each with a very long and slender, slightly curving spine and a small, triangular ventral

one; marginal rim raised, setiferous tubercles well developed, on small ‘ravelins’.

Limbus. With broadly rounded lobes, separated by short stretches ( ca. !4-!A lobe width) of straight

margin; lobes ca. as broad as long; deeply striate on external surface. Surface between lobes forming

shallow concavity behind straight margin.

Tarsal setation (Fig. 5A). Normal.

Gonopod coxa (Figs 9, 17). About 3 ’A x as long as wide. Lateral margin basally straight, then curving

slightly laterad and forming lateral triangular process (Ip) before continuing to rounded coxal tip;

cucullus (cu) hence approximately triangular in outline. Metaplical flange (inf) ending in approximately

equilateral triangular process; metaplical mesal margin straight up to level of metaplical shelf (ms),

ms simple, regularly rounded, projecting mesad as well as posteriad, continuing anterior-mesally in

metaplical shelf-spine (mss) without a marked transition, mss short to medium long, curving first

anteriad, then mesad, then posteriad, almost only in horizontal plane.

Gonopod telopodite (Fig. 17). Solenomere without spines or other secondary branches. Telomere

distally with two branches, both curving 180° anteriad: Anterior lamella (al) smaller than posterior

lamella (pi), both with several lobes with partly serrate-laciniate edges. Due to 180° curvature, distal

part of pi dominating the anterior view, forming a roughly ellipsoid structure with the long axis oriented

basal-apical.

Distribution and habitat

Known only from Mwa nih ana FR. Altitudinal range: 1800-1850 m asl. Habitat: montane rain forest.

Coexisting species

None known.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Chaleponcus basiliscus sp. nov.

um:lsid:zoobank.org:act:AFC2BE4A-F296-4A7E-A396-0C99903FBBBE

Figs 9, 18

Diagnosis

Large. Differing from all other species in the group by the double lateral coxal process (Fig. 9).

Etymology

The name is a noun in apposition and refers to the (remote) resemblance of the cucullus profile to the

reptile Basiliscus or its mythological namesake.

Material studied (total: 4 SS)

Holotype

TANZANIA: $, Iringa Region, Iringa District, Udzungwa Mts, West Kilombero FR, Udekwa village,

Nyambanito Mt., Ukami Forest, 07° 42’49” S, 36°25’15” E, Jul.-Nov. 1994. D. Moyer leg. (ZMUC).

Paratypes

TANZANIA: 3 $$, data as holotype (ZMUC).

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, West Kilombero FR, Udekwa village,

Nyambanito Mt, Ukami Forest, 07° 42’49” S, 36°25 , 15” E.

Description (male)

Diameter. 2.9-3.2 mm, 51-52 podous rings.

Colour. After 20 years in alcohol with traces of a longitudinal light stripe flanked by paramedian darker

bands.

Anal valves. Each with a long, curved dorsal spine and a tiny ventral one.

Limbus. With triangular lobes; lobes slightly longer than broad, striate on outer surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 18). About 3 A x as long as width of shaft. Lateral margin straight up to level of

metaplical shelf (ms), then curving strongly laterad and forming two triangular lateral processes, Ipl and

lp2; distal margin regularly convex; cucullus (cu) mesally ending in process with two blunt-triangular

processes on basal side. Metaplical shelf (ms) regularly rounded, mainly projecting mesad. Metaplical

shelf-spine (mss) long. The position of mss on Fig. 18 is abnormal or an artefact - in the left gonopod of

the specimen in question, and also in the other specimens of C. basiliscus sp. nov., mss is directed first

distad, then regularly curving mesad-basad, as in many other species.

Gonopod telopodite (Fig. 18). Solenomere with a very long, almost straight proximal spine (ps); the

erect position of ps may be abnormal or an artefact - in other gonopods studied the spine is directed

mesad. Telomere distally divided into two relatively small, only slightly lobed lamellae with almost

smooth margins.

Distribution and habitat

Known only from Nyambanito Mt. in West Kilombero Scarp FR. No altitude or habitat information.

European Journal of Taxonomy 100: 1-75 (2014)

Coexisting species

C. ibis sp. nov. was found in the same sample as C. basiliscus sp. nov. In addition, C. circumvallatus sp.

nov., C. gracilior sp. nov., C. netus sp. nov. and C. tintin sp. nov. were found in West Kilombero Scarp FR.

Fig. 18. Chaleponcus basiliscus sp. nov., right gonopod. A. Anterior view. B. Posterior view. C. Mesal

view, cu = cucullus, Ipl and lp2 = lateral coxal processes, ms = metaplical shelf, mss = metaplical shelf-

spine, ps = proximal spine of solenomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss) is

long and projects ± mesad

Chaleponcus krai sp. nov.

urn:lsid:zoobank.org:act:751A38Cl-6F20-4375-A119-79C08025F6C5

Figs 4, 9, 19

Diagnosis

Farge. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine long, stout. Telomere

with characteristically shaped anterior lamella divided into a horizontal lobe with deeply laciniate edges

and a long, straight, distal spine-like part.

Etymology

The name honours Otto Kraus in recognition of his extremely valuable monographs on Odontopygidae;

the name is constructed on a pretended Fatin origin of his surname.

Material studied (total: 6 S3)

Holotype

TANZANIA: 3, Iringa Region, Iringa District, New Dabaga/Ulangambi FR, 08°05’ 37.9” S, 35°54’

05.6” E, fallow/cultivation, 1900-1920 m asl, plot FP, casual, 26-28 Nov. 2000, Frontier Tanzania

UMPS leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District: 2 33, data as holotype (ZMUC); 3 S3, Kalimbazi

Mtn, near Mazombe, 2000-2100 m asl, forest floor, Jan. 1984, Jan Kielland (VMNH).

Type locality

TANZANIA: Iringa Region, Iringa District, New Dabaga/Ulangambi FR, 08°05’ 37.9” S, 35°54’ 05.6”

E, fallow/cultivation, 1900-1920 m asl.

Description (male)

Diameter. 3.0-3.2 mm, 51 podous rings.

Colour. After 12 years in alcohol faded, with traces of pale dorsal band, bordered by dark paramedian

bands.

Anal valves. Each with a long, slightly curving dorsal spine and a well-developed ventral one; marginal

rim raised, setiferous tubercles on ‘ravelins’.

Fimbus (Fig. 4G). With low, apically truncated and strongly serrated lobes; lobes less than half as long

as wide, indistinctly striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 19). Almost 4 x as long as wide. Fateral margin regularly convex from base to

tip; cucullus (cu) small, triangular in outline. Metaplical flange ending in long, triangular process ( mfp ),

pointing distad. Metaplical shelf (ms) regularly rounded, projecting posteriad as well as mesad just

distal to arculus. Metaplical shelf-spine (mss) originating from anterior-mesal end of ms , massive, long,

regularly curved, first anteriad, then disto-mesad, in situ crossing over with opposite mss.

European Journal of Taxonomy 100: 1-75 (2014)

Gonopod telopodite (Fig. 19). Solenomere with very long, straight proximal spine {ps), distally with

three branches:

• an anterior lamella (al) divided into a horizontal lobe with deeply laciniate edges and a long, straight,

distal spine-like part.

Fig. 19. Chaleponcus krai sp. nov., gonopods. Specimens from New Dabaga-Ulangambi (A) and

Kalimbazi Mtn (B-D). A. Left gonopod, anterior view. B-D. Right gonopod. B. Mesal-dorsal view. C.

Telopodite, mesal view. D. Posterior view, al = anterior lobe of telomere, cu = cucullus, il = intermediate

lobe of telomere, mfp = metaplical flange process, mss = metaplical shelf-spine, pi = posterior lobe of

telomere, ps = proximal spine of solenomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

• a large, simple intermediate lamella (z7), projecting distad and closely appressed to spine-like part

of al,

• a small, inconspicuous, rounded posterior lamella (pi).

Distribution and habitat

Known from New Dabaga-Ulangambi FR and from Kalimbazi Mtn, near Mazombe. Altitudinal range:

1900-2100 m asl. Habitat: fallow/cultivation; forest floor.

Coexisting species

In New Dabaga-Ulangambi FR C. gracilior sp. nov. was found in the same sample as C. krai sp. nov. In

addition, C. vilici sp. nov., C. teres sp. nov., C. termini sp. nov., C. netus sp. nov., C. nectarinia sp. nov.,

C. malleolus sp. nov., C. mwabvui sp. nov., C. dabagaensis and C. vandenspiegeli sp. nov. occur in New

Dabaga/Ulangambi FR. C. gracilior sp. nov. also occurs on Kalimbazi Mtn.

Note

The anterior telomeral lamella of C. krai sp. nov. is remarkably similar to the posterior lamella in C.

nectarinia sp. nov., cf note in the general description of the C. dabagaensis group.

Chaleponcus nectarinia sp. nov.

urn:lsid:zoobank.org:act:CA316ECB-0AD5-450C-B2A8-ED38EEE4DA87

Figs 6, 9, 20

Diagnosis

Medium-sized. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine long. Telomere

with characteristically shaped posterior lamella divided into a horizontal lobe with spinose edges and a

long, straight, distal spine-like part.

Etymology

The name (noun in apposition) refers to the somewhat sunbird (Nectarinia)-\ikQ profile of the gonopod

coxal tip.

Material studied (total: 6 <$<$)

Holotype

TANZANIA: 3, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°03’

34.9” S, 35°54’ 41.9” E, montane, 1955 m asl, plot 24, casual, 2 Nov. 2000, Frontier Tanzania leg.

(ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, all

collected by Frontier Tanzania, all in ZMUC: 2 33, 08°03’ 34.9” S, 35°54’ 41.9” E, montane, 1955 m

asl, plot 24, casual, 2 Nov. 2000; 1 3, 08°03’ 39.9” S, 35°54’ 41.9” E, montane, 1945 m asl, plot 24,

29 Oct. 2000; 1 3, 08°04’ 05.7” S, 35°54’ 13.8” E, montane, 1940 m asl, plot 1, sample 1 of 3, 16 Oct.

2000; 1 3, 08°04’ 05.7” S, 35°54’ 18.8” E, montane, 1930-1950 m asl, plot 1, casual, 16-19 Oct. 2000.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°03’ 34.9”

S, 35°54’ 41.9” E, montane, 1955 m asl.

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 20. Chaleponcus nectarinia sp. nov., gonopods. A. Right gonopod, posterior (-lateral) view. B. Right

gonopod, anterior view. C. Tip of posterior lamella of telomere. D. Left gonopod, mesal (-posterior)

view. E. Right gonopod, mesal view, al = anterior lobe of telomere, cu = cucullus, mfp = metaplical

flange process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior lobe of telomere, ps =

proximal spine of solenomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Description (male)

Diameter. 2.3-2.4 mm, 48—49 podous rings.

Colour. After 13 years in alcohol still with a broad light middorsal band, dorsolaterally dark brownish,

laterally brownish-yellow.

Anal valves (Fig. 6A, E). Each with a long, slightly curving dorsal spine and a well-developed, triangular

ventral one; marginal rim raised, setiferous tubercles well developed, on ‘ravelins’.

Fimbus. With long, triangular, apically rounded lobes; lobes ca. twice as long as broad, striate on external

surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 20). About 4 x as long as wide. Fateral margin slightly convex, continuous with

convex apical margin; cucullus (cu) hence regularly rounded; proplical lobe in anterior view hidden

by projecting baso-mesal corner of cucullus. Metaplical flange ending in approximately equilateral

triangular process (mfp); metaplical mesal margin basally shallowly concave, with semicircular incision

at level of arculus. Metaplical shelf (ms) projecting mainly mesad, with approximately right-angled

meso-posterior corner. Metaplical shelf-spine (mss) medium length, arising from distal surface of ms,

first directed apicad, then curving mesad almost in one plane, tip pointing meso-basad.

Gonopod telopodite (Fig. 20). Solenomere with a long, gently curved, proximal spine (ps) originating

next to a wrinkled area. Telomere distally with two branches, both curving anteriad:

• a broad anterior lamella (al) with approximately smooth edges,

• a slender posterior lamella (pi) dividing into a long apicad spine and an anteriad lobe with spinose

edge.

Distribution and habitat

Known only from New Dabaga/Ulangambi FR. Altitudinal range: 1930-1955 m asl. Habitat: montane

forest.

Coexisting species

C. malleolus sp. nov., C. netus sp. nov., C. teres sp. nov., C. termini sp. nov., and C. vilici sp. nov. were

found in the same samples as C. nectarinia sp. nov. In addition, C. dabagaensis, C. gracilior sp. nov., C.

krai sp. nov., C. mwabvui sp. nov. and C. vandenspiegeli sp. nov. occur in New Dabaga/Ulangambi FR.

Note

The posterior telomeral lamella of C. nectarinia sp. nov. is remarkably similar to the anterior lamella in

C. krai sp. nov., cf. note in the general description of the C. dabagaensis group.

Chaleponcus circumvallatus sp. nov.

um:lsid:zoobank.org:act:67292AA6-2D3C-4DE4-BCF0-55719766C6E7

Figs 4-6, 9, 21

Diagnosis

Farge. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine long. Telomere with

a smooth anterior lamella and a posterior lamella ending in two long spines. Claws of walking legs

surrounded by ‘palisade’ of setae, a character shared only with C. ibis sp. nov.

European Journal of Taxonomy 100: 1-75 (2014)

Etymology

The name is a Latin adjective meaning ’’surrounded by palisade” and refers to the ‘palisade’ of setae

surrounding the claws.

Material studied (total: 2 ff)

Holotype

TANZANIA: S, Tanzania, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village,

1600-1650 m, 8-12 Nov. 1984, montane rain forest, N. Scharff leg. (ZMUC).

Paratype

TANZANIA: 1 S, Iringa Region, Iringa District, 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 UMPS leg.

(ZMUC).

Type locality

TANZANIA: Tanzania, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village,

1600-1650 m, montane rain forest.

Description (male)

Diameter. 2.8 mm, 48 podous rings.

Colour. After 14 years in alcohol uniform straw yellow, metazonites slightly darker, no traces of a

dorsal light band.

Anal valves (Fig. 6B). Each with a very long and stout, slightly curving dorsal spine and a very long

and slender ventral one; marginal rim raised; setiferous tubercles well developed, on small ‘ravelins’.

Limbus (Fig. 4J). Almost rectilinear, with just the slightest indication of lobes.

Tarsal setation (Fig. 5 B-C). Unusual: claws surrounded and almost hidden by numerous stiff setae,

accessory claw much longer than claw.

Gonopod coxa (Figs 9, 21). Stout, ca. 3 x as long as wide. Lateral margin slightly convex in basal %,

then straight, then continuing in smooth curve in apical margin; cucullus (cu) hence regularly rounded.

Metaplical flange ending in long triangular process ( mfp); metaplica posterior-mesally with large

triangular process ( mp ), delimiting semicircular sinus together with base of metaplical shelf. Metaplical

shelf (ms) regularly rounded, projecting mesad as well as posteriad. Metaplical shelf-spine (mss)

originating from anterior-mesal end of ms , massive, long, regularly curved, first anteriad, then disto-

mesad.

Gonopod telopodite (Fig. 21). Solenomere with a very long, straight, mesad proximal spine (ps ).

Telomere distally with two branches, both curving anteriad:

• a rounded anterior lamella (al) with small, rectangular accessory lamella (al ’) on its mesal surface,

• a posterior lamella (pi) ending in an apicad and a basad-anteriad spine.

Distribution and habitat

Known from West Kilombero Scarp FR and Udzungwa Scarp FR. Altitudinal range 1390-1650 m asl.

Habitat: montane forest.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Coexisting species

C. hamerae sp. nov. and C. nikolajscharffi sp. nov. also occur in Udzungwa Scarp FR. In West Kilombero

Scarp FR C. netus sp. nov. was found in the same sample as C. circumvallatus sp. nov. In addition, C.

basiliscus sp. nov., C. gracilior sp. nov., C. ibis sp. nov. and C. tintin sp. nov. occur in West Kilombero

Scarp FR.

Fig. 21. Chaleponcus circumvallatus sp. nov., left gonopod. A. Anterior view. B. Mesal-anterior view.

C. Posterior view. D. Mesal-posterior view, al = anterior lobe of telomere, aV = accessory lamella, cu =

cucullus, mfp = metaplical flange process, mp = mesal metaplical process, ms = metaplical shelf, mss =

metaplical shelf-spine,/?/ = posterior lobe of telomere,^ = proximal spine of solenomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Notes

Resembles the smaller C. nectarinia sp. nov. and the similar-sized C. krai sp. nov. in gonopod

morphology, including in the possession of a telomeral lobe with a long spine. Differs from these species

in the detailed shape of this lobe. Differs from all other species in the C. dabagaensis group except C.

ibis sp. nov. by the peculiar distal setation of the walking leg tarsi. Although females are generally not

considered in this study, it deserves mention that a female specimen found together with the male C.

circumvallatus sp. nov. from West Kilombero Scarp FR and sharing its size and general morphology,

also has the strange tarsal setation. The function of this is uncertain; maybe it is an adaptation for

walking on soft substrates (I.B. Enghoff pers. comm.).

Chaleponcus ibis sp. nov.

urn:lsid:zoobank.org:act:E6764255-8F2F-4B2D-8F3C-26F40FA01EB0

Figs 9, 22

Diagnosis

Large. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine very long. Telomere

with a rugged anterior lamella and posterior lamella ending in short, parallel, elongate lobes. Claws of

walking legs surrounded by ‘palisade’ of setae, a character shared only with C. circumvallatus sp. nov.

Etymology

The name is a noun in apposition and refers to the somewhat ibis bill-like metaplical shelf-spine.

Material studied (total: 1 S)

Holotype

TANZANIA: S, Iringa Region, Iringa District, Udzungwa Mts, West Kilombero FR, Udekwa village,

Nyambanito Mt., Ukami Forest, 07° 42’49” S, 36°25’ 15” E, Jul.-Nov. 1994. D. Moyer leg. (ZMUC).

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, West Kilombero FR, Udekwa village,

Nyambanito Mt., Ukami Forest, 07° 42’49” S, 36°25’15” E.

Description (male)

Diameter. 3.5 mm, 54 podous rings (the largest of all species in the group).

Colour. After 20 years in alcohol ground colour greyish, posterior part of metazonites yellow-brownish;

traces of a broad dorsal dark stripe (most other species have traces of a light stripe).

Anal valves. Each with a strong, apparently broken dorsal and smaller, but well-developed ventral

spine. Margin raised, setiferous tubercles on ravelins.

Limbus. With triangular lobes; lobes ca. as long as wide, margin micro-dentate, outer surface striate.

Tarsal setation. Unusual: claws surrounded and almost hidden by numerous stiff setae, accessory claw

much longer than claw (as in C. circumvallatus sp. nov., Fig. 5B, C).

Gonopod coxa (Figs 9, 22). Stout, ca. 3 x as long as wide. Lateral margin irregularly convex, continuing

in smooth curve in apical margin; cucullus (cu) hence regularly rounded. Metaplica posterio-mesally

with triangular process imp). Metaplical shelf {ms) relatively small, giving rise to very long, slightly

curved metaplical shelf-spine (mss).

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Gonopod telopodite (Fig. 22). Solenomere with a very long, straight, mesad proximal spine (ps).

Telomere distally with two branches, both curving anteriad:

• an irregular shaped anterior lamella ( al) with coarsely dentate proximal part and a smooth,

subrectangular distal part (which maybe corresponds to the accessory lobe, al \ in C. circumvallatus

sp. nov.),

• a posterior lamella (pi) ending in two short, parallel, elongate lobes.

Fig. 22. Chaleponcus ibis sp. nov., left gonopod. A. Posterior view. B. Anterior view. C. Telopodite,

mesal-anterior view, al = anterior lobe of telomere, al ’ = accessory lamella?, cu = cucullus, mp = mesal

metaplical process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior lobe of telomere,

ps = proximal spine of solenomere. Scales 0.1 mm

European Journal of Taxonomy 100: 1-75 (2014)

Distribution and habitat

Known only from Nyambanito Mt. in West Kilombero Scarp FR. No altitude or habitat information.

Coexisting species

C. basiliscus sp. nov. was found in the same sample as C. ibis sp. nov. In addition, C. circumvallatus sp.

nov., C. gracilior sp. nov., C. netus sp. nov. and C. tintin sp. nov. were found in West Kilombero Scarp

FR.

Notes

Resembles the almost similar-sized C. circumvallatus sp. nov. both in the unusual tarsal setation and

in gonopod structure. C. ibis sp. nov. differs from C. circumvallatus sp. nov., i.a., in having distinct

lobes on the limbus, in having the metaplical shelf-spine (mss) much longer, in having the proximal

part of the anterior telomeral lobe (al) coarsely dentate, and in having two relatively short, parallel

elongate lobes on the posterior telomeral lamella, as opposed to the two very long, diverging spines in

C. circumvallatus sp. nov.

Chaleponcus vcindenspiegeli sp. nov.

urn:lsid:zoobank.org:act:A2C22BA2-915D-47DC-A49C-9EE26F52F8D9

Figs 9, 23

Diagnosis

Medium-sized. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine long. Telomere

with an anterior lamella ending in two short, parallel processes and a posterior lamella with serrate

margin.

Etymology

The name honours Didier VandenSpiegel in recognition of his taxonomic work on Afrotropical

millipedes.

Material studied (total: 1 f )

Holotype

TANZANIA: Iringa Region, Iringa District, New Dabaga/Ulangambi FR, 08°05’ 34.5” S, 35°55’

31.5” E, montane, 1800-1900 m asl, plot 16, casual, 15-16 Nov. 2000, Frontier Tanzania leg. (ZMUC).

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°05’ 34.5”

S, 35°55’ 31.5” E, 1800-1900 m asl.

Description (male)

Diameter. 2.0 mm, 44 podous rings.

Colour. After 14 years in alcohol generally straw-yellow, with a broad, light middorsal stripe formed by

an hourglass-shaped spot on each body ring; stripe flanked by dark paramedian zones.

Anal valves. Each with a small dorsal and a very small ventral denticle; mesal margin raised, setiferous

tubercles not on ravelins.

Limbus. With slender triangular lobes; lobes ca. twice as long as broad.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 23). About 3A x as long as broad. Metaplical flange ending in rounded,

projecting lobe ( mfp ); metaplica with large rounded process (mp) just basal to arculus. Metaplical shelf

(ms) relatively small. Metaplical shelf-spine (mss) medium-long, directed mesad, bent at ca. 120° at

midlength.

Fig. 23. Chaleponcus vandenspiegeli sp. nov., left gonopod. A. Anterior view. B. Mesal view. C. Tip

of telomere and part of solenomere, ventral view. D. Posterior view, al = anterior lamella of telomere,

alpl and alp2 = processes from al, mfp = metaplical flange process, mp = mesal metaplical process, ms

= metaplical shelf, mss = metaplical shelf-spine, pi = posterior lamella of telomere , ps = proximal spine

of solenomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Gonopod telopodite (Fig. 23). Solenomere with a medium-long proximal spine (ps). Telomere distally

with two branches which both bend anteriad:

• a relatively simple anterior lamella (al) with a number of lobes etc. along its margin, including two

short, parallel processes (< alpl , alp2),

• a strongly convoluted posterior lamella (pi) which because of its bending over is largely visible in

anterior view; margin of pi strongly serrate.

Distribution and habitat

Known only from Udzungwa Mts, New Dabaga/Ulangambi FR. Altitudinal range: 1800-1900 m asl.

Habitat: montane forest.

Coexisting species

C. gracilior sp. nov., C. netus sp. nov. and C. termini sp. nov. were found in the same sample as C.

vandenspiegeli sp. nov. In addition, C. dabagaensis, C. krai sp. nov., C. malleolus sp. nov., C. mwabvui

sp. nov., C. nectarinia sp. nov., C. teres sp. nov. and C. vilici sp. nov. occur in New Dabaga/Ulangambi

FR.

Note

The gonopods of C. vandenspiegeli sp. nov. somewhat resemble those of the much bigger species C.

krai sp. nov. The two processes of the anterior telopodal lamella in C. vandenspiegeli sp. nov. are in

almost the same position in relation to the rest of the gonopod as the two parallel lobes on the posterior

telopodal lamella in C. ibis sp. nov.

Species without an obvious lateral coxal process in which the metaplical shelf-spine (mss) is

not long and projecting ± mesad

(C. vilici sp. nov. has the lateral side of the coxa somewhat angled which reminds of a small lateral

process)

Chaleponcus vilici sp. nov.

urn:lsid:zoobank.org:act:F44D3A63-lE8A-4D6E-AE7D-6BC61723D34F

Figs 8-9, 24

Diagnosis

Medium-sized. Differs from all other species of the C. dabagaensis group by the two-levelled metaplical

shelf and the course of the metaplical shelf-spine, which fits between the two levels of the shelf. Gonopod

coxa (Fig. 9) without a lateral process; metaplical shelf-spine long.

Etymology

The name honours Richard L. Hoffman (1927-2012) global grand-master of diplopodology through

much of the last half of the previous, as well as the first decade of the present century. Vilici is the

genitive case of the Latin noun vilicus, meaning ‘steward’, which is also the original meaning of the

name Hoffman.

Material studied (total: 37 <$<$)

Holotype

TANZANIA: Iringa Region, Iringa District, New Dabaga/Ulangambi FR, 08°04’ 05.7” S, 35°54’

18.8” E, montane, 1930-1950 m asl, plot 1, casual, 16-19 Oct. 2000, Frontier Tanzania leg. (ZMUC).

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 24. Chaleponcus vilici sp. nov., left gonopod. A. Posterior view. B. Mesal view. C. Mesal-anterior

view. D. Close-up of metaplical shelf-spine, mesal view. E. Anterior (and slightly apical/ventral) view.

F. Telopodite, distal view, showing solenomere and the two distal telomere branches, al = anterior lamella

of telomere, cu = cucullus, la = lateral angle of coxa, mf= metaplical flange, mp = mesal metaplical

process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior lamella of telomere, slm =

solenomere. Scales: 0.1 mm (A—D), 0.01 mm (E-F).

European Journal of Taxonomy 100: 1-75 (2014)

Paratypes

All from TANZANIA, Iringa Region, Iringa District, New Dabaga/Ulangambi FR, all collected by

Frontier Tanzania, all in ZMUC: 23 SS, 08°04’ 05.7” S, 35°54’ 18.8” E, montane, 1930-1950 m asl,

plot 1, casual, 16-19 Oct. 2000; 3 $$, 08°00’ 26.6” S, 35°56’ 06.1” E, montane forest, 1910 m asl, plot

Kinyonga, casual, 24 Oct. 2000; 1 $, 08°00’ 26.5” S, 35°56’ 06.1” E, scrub/thicket/bush, 1908 m asl,

plot Kinyonga, 19 Oct. 2000; 1 $, 08°04’ 05.7” S, 35°54’ 13.8” E, montane, 1940 m asl, plot 1, sample

1 of 3, 16 Oct. 2000; 7 $$, 08°03’34.9” S, 35°54’ 41.9” E, montane, 1955 m asl, plot 24, casual, 2 Nov.

2000; 1 $, 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930 m asl, plot 6, sample 3 of 3, 27 Oct. 2000.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°04’ 05.7”

S, 35°54’ 18.8” E, montane, 1930-1950 m asl.

Description (male)

Diameter. 2.0-2.1 mm, 46^19 podous rings.

Colour. After 12 years in alcohol pale brown below ozopores, blackish above, dorsally with a light band

with a thin dark midline, telson blackish.

Anal valves. Each with a long, slightly curving dorsal spine, no ventral spine; marginal rim raised,

setiferous tubercles on ‘ravelins’.

Limbus. With long, triangular, apically rounded lobes, ca. Wi times longer than wide, lobes striate on

external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 24). About 3 x as long as wide. Lateral margin shallowly S-shaped: slightly

convex in middle, then slightly concave, subdistally with a protruding angle (la), apical margin starting

from la oblique, shallowly sinuous; cucullus (cii) hence triangular, apically with small mesad process.

Metaplical flange (mf) ending in blunt, rounded angle, no process; metaplical mesal margin with 3

processes: a rounded, semicircular process (mp) basal to arculus and two processes, formed by metaplical

shelf, distal to arculus. Metaplical shelf (ms) massive, with two levels separated by concavity into which

tip of metaplical shelf-spine (mss) fits, mss long, slender, originating from distal surface of ms, from

there first projecting distad, then turning posteriad at right angles and curving smoothly basad, and then

anteriad, fitting between two levels of ms, in mesal view (Fig. 24D) almost forming a letter D (mss

frequently broken).

Gonopod telopodite (Figs 8B, 24). Solenomere (slm) with a long, straight proximal spine ( ps ). Telomere

distally with two relatively small branches, folded around each other somewhat like a pair of one¬

fingered hands, the ‘finger’ of the anterior lamella (al) having smooth edges, that of the slenderer

posterior lamella (pi) having jagged edges.

Distribution and habitat

Known only from Udzungwa Mts, New Dabaga/Ulangambi FR. Altitudinal range: 1908-1955 m asl.

Coexisting species

C. gracilior sp. nov., C. malleolus sp. nov., C. nectarinia sp. nov., C. netus sp. nov., C. teres sp. nov. and

C. termini sp. nov. were found in the same samples as C. vilici sp. nov. In addition, C. dabagaensis, C.

krai sp. nov., C. mwabvui sp. nov. and C. vandenspiegeli sp. nov. occur in New Dabaga/Ulangambi FR.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Chaleponcus teres sp. nov.

urn:lsid:zoobank.org:act:F4EB8A22-2F19-438F-8C65-DB6091867337

Figs 4, 6, 9, 25

Diagnosis

Medium-sized. Differs from all other group members, except C. malleolus sp. nov., by the absence of

a dorsal spine on each anal valve. Further characterized by the subquadratic outline of the cucullus and

the structure of the telomere, especially the presence of a slender, sinuous, spine-like branch. Gonopod

coxa (Fig. 9) without a lateral process; metaplical shelf-spine long.

Etymology

The name is a Fatin adjective meaning “smooth” and refers to the spineless anal valves.

Material studied (total: 29 SS)

Holotype

TANZANIA: c?, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°04’

05.7” S, 35°54’ 18.8” E, montane, 1930-1950 m asl, plot 1, casual, 16-19 Oct. 2000, Frontier Tanzania

leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, all

collected by Frontier Tanzania, all in ZMUC: 27 08°04’ 05.7” S, 35°54’ 18.8” E, montane, 1930-

1950 m asl, plot 1, casual, 16-19 Oct. 2000; 1 08°04’ 05.7” S, 35°54’ 13.8” E, montane, 1940 m asl,

plot 1, sample 3 of 3, 19 Oct. 2000.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°04’ 05.7”

S, 35°54’ 18.8” E, montane, 1930-1950 m asl.

Description (male)

Diameter. 1.9-2.2 mm, 45—47 podous rings.

Colour. After 12 years in alcohol yellowish, with dark spots at ozopores and paramedian dark dorsal

bands flanking a mid-dorsal light stripe; paramedian dark colouration sometimes extending down flanks

on metazonites, giving the body a ringed appearance; body rings 6 and 7 darker than the others.

Anal valves (Fig. 6D). Entirely without dorsal and ventral spines; marginal rim not raised, setae not on

tubercles.

Fimbus (Fig. 4H). With low, broad, rounded lobes, separated by parabolic sinuses; each lobe ca. 4 times

broader than long, margin densely and finely serrate, lobes indistinctly striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 25). About 3Vi x as long as wide. Fateral margin straight in basal %, then

shallowly concave, angle between lateral and apical margins 90°, sharp; cucullus (cu) hence subquadratic.

Metaplical flange ( mf) smoothly rounded, no angle, no process; metaplical mesal margin straight up to

level of arculus. Metaplical shelf (ms) forming subrectangular process just distal to arculus, followed by

very deep incision (in) separating cucullus from basal part of metaplica; ms proximally stout, continuing

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 25. Chaleponcus teres sp. nov., gonopods. A. Gonopods in situ in seventh body ring, anterior view.

B. Left gonopod, mesal view. C. Right gonopod, base of solenomere and telomere, ventral view. D. Right

gonopod, posterior view. E. Left gonopod, anterior(-lateral-ventral) view. F. Right telopodite, mesal-

anterior-ventral view. G. Right telopodite, mesal-anterior view, all, al2 = branches of apical telomeral

lamella, bal, bal’ = basal lamella of telomere, cu = cucullus, in = metaplical incision, mf= metaplical

flange, ms = metaplical shelf mss = metaplical shelf-spine, ps = proximal spine of solenomere, slm =

solenomere, spl = spine-like lamella of telomere, st, st ’ = main stem of telomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

in metaplical shelf-spine (mss) with no clear demarcation, mss medium long, originating from anterior-

distal surface of ms , first projecting distad, then curving mesad, and then slightly basad.

Gonopod telopodite (Fig. 25). Solenomere with a small, inconspicuous, irregular proximal spine (ps).

Telomere without a separate proximal lobe but posterior margin of proximal part of main stem ( st ’) lying

closely against base of solenomere (slm). Telomere distally with three branches:

• A large, roughly rectangular basal lamella ( bal ) carrying a secondary lamella ( bal ’) on its lateral

side,

• A distal spine-like branch, slender and sinuous (spl),

• An irregular apical lamella dividing in two branches, proximal branch (all) with smooth margins,

the distal branch (al2) with irregularly serrated margin, all and bal in situ curving around opposite

coxa (Fig. 25A).

Distribution and habitat

Known only from Udzungwa Mts, New Dabaga/Ulangambi FR. Altitudinal range: 1930-1950 m asl.

Habitat: montane forest.

Coexisting species

C. termini sp. nov. and C. vilici sp. nov. were found in the same samples as C. teres sp. nov. In

addition, C. dabagaensis, C. gracilior sp. nov., C. krai sp. nov., C. malleolus sp. nov., C. mwabvui sp.

nov., C. nectarinia sp. nov., C. netus sp. nov. and C. vandenspiegeli sp. nov. occur in New Dabaga/

Ulangambi FR.

Notes

The gonopods somewhat resemble those of C. hamerae sp. nov.

Chaleponcus hamerae sp. nov.

urn:lsid:zoobank.org:act:E27E3FB0-22E6-46EF-81CB-447DA44E3090

Figs 9, 26

Diagnosis

Medium-sized. Characterized by the sub-rectangular outline of the cucullus, resembling C. teres sp. nov.

in this character, but differing by having a dorsal spine on each anal valve. Telomeral lamellae largely

smooth. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-spine long.

Etymology

The name honours Michelle Hamer in recognition of her work on the taxonomy and conservation of

African millipedes.

Material studied (total: 9 33 )

Holotype

TANZANIA: 3 , Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village, 1600-1650

m, 8-13 Nov. 1984, pitfall traps, montane rain forest, N. Scharff leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, all in ZMUC: 4 S3, above

Chita village, 1600-1650 m, 8-13 Nov. 1984, pitfall traps, montane rain forest, N. Scharff leg; 4 33 ,

11 km SE of Masisiwe village, Kihanga Stream, 1800 m, 08°22’ 05.7” S, 35°58’ 41.6” E, 17-27 May

1997. ZMUC & SI Exp.

European Journal of Taxonomy 100: 1-75 (2014)

Type locality

TANZANIA: Iringa Region, Udzungwa Mts, Udzungwa Scarp FR, above Chita village, 1600-1650 m,

rain forest.

Description (male)

Diameter. 2.1-2.3 mm, 46—47 podous rings.

Colour. After 30 years in alcohol (brownish) yellow, traces of broad light dorsal stripe flanked by

dorsolateral dark stripes, narrow lateral dark line at ozopore level.

Anal valves. Each with a small dorsal spine and a very small ventral spine; marginal rim raised,

setiferous tubercles on ravelins.

Limbus. With broadly rounded to spatulate, apically finely serrate lobes; lobes as broad as long or slightly

broader, separated by rounded incisions much narrower than width of lobes, or by short stretches of

straight, finely serrulate margin.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 26). About 3V£-4 x as long as wide. Lateral margin basally slightly sinuous,

basally convex, ca. at level of proplical lobe (prl) concave and then again convex, meeting oblique,

straight apical margin under blunt angle; cucullus (cu) hence subrectangular in outline. Metaplical

flange ( mf) ending in blunt, rounded angle, no process; metaplical mesal margin slightly concave up to

level of prl , distally tucked in, on posterior side sinuous: first concave, then convex, then concave again

up to level of metaplical shelf (ms), ms very prominently projecting mesad but not notably posteriad.

Metaplical shelf-spine (mss) short, originating from anterior-mesal part of ms, projecting distad, slightly

curved.

Gonopod telopodite (Fig. 26). Solenomere without a proximal spine. Telomere distally with two distal

branches, both curving anteriad:

• anterior lamella (al) broad, twisted, mostly with smooth edges but with terminal, subrectangular

lobe (al ’) with dentate terminal edge,

• posterior lamella (pi) situated basal to al and rest of telopodite, large, smoothly curved, with smooth

edges and subterminally with long, apical spine (pis).

Distribution and habitat

Known only from Udzungwa Mts, Udzungwa Scarp FR. Altitudinal range: 1600-1800 m asl. Habitat:

montane rain forest.

Coexisting species

C. nikolajscharffi sp. nov. was found in the same samples as C. hamerae sp. nov. In addition, C.

circumvallatus sp. nov. was found in Udzungwa Scarp FR.

Notes

The gonopods quite resemble those of C. teres sp. nov.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 26. Chaleponcus hamerae sp. nov., left gonopod. A. Mesal-anterior view. B. Posterior view. C. Anterior

view. D. Ventral view, al, al' = anterior telomeral lamella, cu = cucullus, mf= metaplical flange, ms =

metaplical shelf, mss = metaplical shelf-spine, pi = posterior telomeral lamella, prl = proplical lobe, pis

= spine on pi. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Chaleponcus termini sp. nov.

um:lsid:zoobank.org:act:C7102566-C098-434C-A419-652D26ED1738

Figs 7-9, 27

Diagnosis

Medium-sized. Characterized by the two-lobed, strongly serrate anterior telomeral lamella. Gonopod

coxa (Fig. 9) without a lateral process; metaplical shelf-spine long.

Etymology

The name is the genitive case of the Fatin noun terminus which means frontier, and the name honours

the NGO Frontier Tanzania.

Material studied (total: 28 S<S)

Holotype

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°04’

5.7” S, 35°54’ 18.8” E, montane, 1930-1950 m asl, plot 1, casual, 16-19 Oct. 2000, Frontier Tanzania

leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, all

collected by Frontier Tanzania, all in ZMUC: 4 08°05’ 34.5” S, 35°55’ 31.5” E, montane, 1800-

1900 m asl, plot 16, casual, 15-16 Nov. 2000; 1 S, 08°06’ 47.4” S, 35°56’ 50.4” E, montane, 1800-1955

m asl, plot MS - plot A, 29 Oct. 2000; 10 SS, 08°05’ 34.5” S, 35°55’ 31.8” E, montane, 1885 m asl,

plot 16, sample 2 of 3, 15 Nov. 2000; 1 S, 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930 m asl, plot 6,

sample 1 of 3, 28 Oct. 2000; 1 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930 m asl, plot 6, sample 2

of 3, 27 Oct. 2000; 2 08°04’ 05.6” S, 35°54’ 20.4” E, montane, 1930 m asl, plot 6, sample 3 of 3, 27

Oct. 2000; 7 $$, 08°04’ 5.7” S, 35°54’ 18.8” E, montane, 1930-1950 m asl, plot 1, casual, 16-19 Oct.

2000; 1 c?, 08°04’ 05.7” S, 35°54’ 13.8” E, montane, 1940 m asl, plot 1, sample 1 of 3, 16 Oct. 2000.

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°04’ 5.7”

S, 35°54’ 18.8” E, montane, 1930-1950 m asl.

Description (male)

Diameter. 1.6-1.9 mm, 41—43 podous rings.

Colour. After 12 years in alcohol yellowish, with dark spots at ozopores and paramedian dark dorsal

bands flanking a mid-dorsal light stripe.

Anal valves. Each with a long, slightly curving dorsal spine and a well-developed, triangular ventral

one; marginal rim raised, no setiferous tubercles.

Limbus. With rounded-triangular, longitudinally furrowed denticles separated by parabolic sinuses,

lobes ca. as long as wide, striate on external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 7, 9, 27). Almost 4 x as long as wide. Lateral margin regularly convex from base

up to just below level of prl , at this place turning slightly laterad and forming blunt, rounded angle with

oblique apical margin; cucullus ( cu ) hence subtriangular, apically rounded. Metaplical flange (mf) ending

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

in long, triangular process pointing anteriad-mesad-distad; metaplical mesal margin straight up to level

of arculus. Metaplical shelf (ms) forming subrectangular-rounded process just distal to arculus, followed

by triangular incision separating cucullus from basal part of metaplica, ms simple, proximally stout,

continuing in metaplical shelf-spine (mss) with no clear demarcation, mss medium long, originating

from anterior-distal surface of ms , first projecting distad, then curving mesad, and then slightly basad.

Gonopod telopodite (Figs 8A, 27). Solenomere with a small, irregular proximal spine (ps). Telomere

distally with two branches:

• a large anterior lamella dividing into two triangular lobes (all and all) with strongly laciniate

margins; basalmost lobe (all) folded laterad and in situ visible in front of opposite coxa,

• a large posterior lamella (pi) first directed distad then folding back and delimiting an empty space

together with al2.

Fig. 27. Chaleponcus termini sp. nov., gonopods. A. Right gonopod, posterior view. B. Right gonopod,

anterior view. C. Feft telopodite, mesal-ventral view. D. Feft coxite, mesal view. E. Right telopodite,

anterior-ventral view, all, al2 = anterior telomeral lobe, cu = cucullus, mf= metaplical flange, ms =

metaplical shelf, mss = metaplical shelf-spine, pi = posterior telomeral lamella, ps = proximal spine on

solenomere. Scales 0.1 mm.

European Journal of Taxonomy 100: 1-75 (2014)

Distribution and habitat

Known only from Udzungwa Mts, New Dabaga/Ulangambi FR. Altitudinal range: 1800-1955 m asl.

Habitat: montane forest.

Coexisting species

C. gracilior sp. nov., C. mwabvui sp. nov., C. nectarinia sp. nov., C. netus sp. nov., C. teres sp. nov. and

C. vilici sp. nov. were found in the same samples as C. termini sp. nov. In addition, C. dabagaensis, C.

krai sp. nov., C. malleolus sp. nov. and C. vandenspiegeli sp. nov. occur in New Dabaga/Ulangambi FR.

Chaleponcus gracilior sp. nov.

um:lsid:zoobank.org:act:85282955-A517-4E67-8848-F2C3C880DEC6

Figs 4, 9, 28

Diagnosis

Slender, but with more podous rings than other small species. Easily recognizable by the subrectangular

cucullus profile, by the conspicuous ‘inflated’ distal telomeral process and by the unique ‘string-of-

beads’-like appearance of the limbus. Gonopod coxa (Fig. 9) without a lateral process; metaplical shelf-

spine short.

Etymology

The name is a Latin adjective meaning “more slender” and refers to the small body diameter in relation

to the number of body rings.

Material studied (total: 84 33 )

Holotype

TANZANIA: 3 , Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°05’

37.9” S, 35°54’ 05.6” E, fallow/cultivation, 1900-1920 m asl, plotFP, casual, 26-28 Nov. 2000, Frontier

Tanzania leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, all

collected by Frontier Tanzania, all in ZMUC: 1 3 , 08°05’34.5” S, 35°55’31.5” E, montane, 1800-1900

m asl, plot 16, casual, 15-16 Nov. 2000; 1 3, 08°06’ 47.4” S, 35°56’ 50.4” E, montane, 1800-1955 m

asl, plot MS - plot A, 29 Oct. 2000; 2 33 , 08°03’ 39.9” S, 35°54’ 05.6” E, fallow/previously disturbed,

1900 m asl, plot FF, sample 2 of 3, 28 Nov. 2000; 18 33 , 08°05’ 37.9” S, 35°54’ 05.6” E, fallow/

cultivation, 1900-1920 m asl, plot FP, casual, 26-28 Nov. 2000; 3 33 , 08°00’ 26.5” S, 35°56’ 06.1”

E, scrub/thicket/bush, 1908 m asl, plot Kinyonga, 19 Oct. 2000; 3 33 , 08°00’ 26.6” S, 35°56’ 06.1” E,

montane forest, 1910 m asl, plot Kinyonga, casual, 24 Oct. 2000; 2 33 , 08°03’ 39.9” S, 35°54’ 05.6”

E, fallow/previously disturbed, 1915 m asl, plot FF, casual, 27 Nov. 2000; 1 3 , 08°00’ 26.6” S, 35°56’

06.1” E, scrub/thicket/bush, 1915 m asl, plot Kinyonga, 18 Oct. 2000; 1 3 , 08°03’ 43.6” S, 35°53’ 54.2”

E, plantation, 1980 m asl, plot pine, sample 1 of 3, 13 Oct. 2000.

Referred non-type material

TANZANIA: 47 33 , Iringa Region, Iringa District, West Kilombero Scarp FR, 07°45’ 34.2” S, 36°26’

37.4” E, (open) woodland, 1510 m asl, plot Acacia, casual, 5-8 Dec. 2000, Frontier Tanzania UMPS leg.

(ZMUC); 1 3 , Udzungwa Mts, 1500 m, Kiranzi-Kitungulu FR, 08°09’S, 35°05’E, forest, Jan. 1996, M.

Andersen, P. Gravlund, A. Jalcobsen leg. (ZMUC); 3 33 , Iringa Region, Iringa District, Kalimbazi Mtn,

2000-2100 m asl, 10 km S of Mazombe, forest floor, Jan Kielland leg. (VMNH).

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Fig. 28. Chaleponcus gracilior sp. nov., gonopods. A. Gonopods in situ in body ring 7, anterior view.

B. Right gonopod, detail of base of solenomere, showing basal side branch consisting of a tiny, spine¬

like part and a larger, irregularly lamellate one. C. Right gonopod, anterior-mesal view. D. Feft gonopod,

posterior view. E. Feft telopodite, posterior-lateral view. F. Right telopodite, anterior view, al = anterior

telomeral lamella, cu = cucullus, dp = distal telomeral process, mf= metaplical flange, ms = metaplical

shelf, mss = metaplical shelf-spine, pb = proximal side branch of solenomere, pi = posterior telomeral

lamella, prl = proplical lobe. Scales: 0.1 mm (A, C-F), 0.05 mm (B).

European Journal of Taxonomy 100: 1-75 (2014)

Type locality

TANZANIA: Iringa Region, Iringa District, New Dabaga/Ulangambi FR, 08°05’ 37.9” S, 35°54’ 05.6”

E, fallow/cultivation, 1900-1920 m asl.

Description (male)

Diameter. 1.5-1.8 mm, 45—49 podous rings.

Colour. After 12 years in alcohol faded, uniform pale brown, no traces of a dorsal stripe.

Anal valves. Each with a long dorsal spine, no ventral spines; marginal rim barely raised, setiferous

tubercles inconspicuous.

Limbus (Fig. 4F). Lobes spatulate, about as long as wide, separated by stretches of straight margin

ca. same length as width of lobe; body ring surface between lobes forming shallow concavity behind

straight margin; lobes deeply striate on external surface. The unique limbus gives the body ring posterior

margins a characteristic ‘string-of-beads’-like appearance, visible even on undissected specimens.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 28). About 4 x as long as wide. Lateral margin almost straight, slightly indented

at level of proplical lobe (prl) and metaplical shelf (ms), forming a ca. 100° angle with straight apical

margin; cucullus (cu) subrectangular. Metaplical flange (mf) ending in a rather sharp 90° angle, no

process; metaplical mesal margin shallowly concave up to level of arculus, then abruptly turning mesad

at right angles at base of metaplical shelf (ms), then turning apicad at right angles, margin from then

on shallowly concave, meeting apical margin under a ca. 80° angle. Metaplical shelf (ms) regularly

rounded, projecting posteriad as well as mesad. Metaplical shelf-spine (mss) originating from anterior-

mesal end of ms, short, directed apicad, slightly and simply curved.

Gonopod telopodite (Fig. 28). Solenomere with a small proximal side branch (pb) originating on distal

surface and consisting of a tiny spine-like part and a larger, irregularly lamellate one. Telomere distally

with three branches:

• an anterior, irregularly triangular, twisted lamella (al) with undulate edges,

• a large, smooth, pointed, slightly curved distal process (dp) with an ‘inflated’ appearance,

• an irregularly shaped posterior lamella (pi).

Distribution and habitat

Known from four areas in the Udzungwa Mts: New Dabaga/Ulangambi FR, West Kilombero Scarp FR,

Kiranzi-Kitungulu FR and Kalimbazi Mtn. Altitudinal range: 1500-2100 m asl. Habitat: montane forest,

fallow/previously disturbed/cultivation, scrub/thicket/bush, plantation, (open) woodland. This species

seems much more euryoecious than any other species in the C. dabagaensis group.

Coexisting species

In line with its comparatively wide distribution, C. gracilior sp. nov. co-occurs with many other species

of the C. dabagaensis group. Thus, C. dabagaensis was found in the same sample as C. gracilior sp.

nov. in Kiranzi-Kitungulu FR, and C. dabagaensis, C. krai sp. nov., C. malleolus sp. nov., C. mwabvui

sp. nov., C. netus sp. nov., C. termini sp. nov. and C. vilici sp. nov. were found in the same samples as C.

gracilior sp. nov. in New Dabaga/Ulangambi FR. In addition, C. krai sp. nov. also occurs on Kalimbazi

Mtn, C. nectarinia sp. nov., C. teres sp. nov., and C. vandenspiegeli sp. nov. also occur in New Dabaga/

Ulangambi FR., and C. basiliscus sp. nov., C. circumvallatus sp. nov., C. ibis sp. nov., C. netus sp. nov.,

and C. tintin sp. nov. also occur in West Kilombero FR.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Chaleponcus mwabvui sp. nov.

urn:lsid:zoobank.org:act:ClBCEEF3-3247-47D9-BA27-663385735829

Figs 6, 9, 29

Diagnosis

Medium-sized. Gonopod coxa (Fig. 9) without a lateral process; cucullus small. Distinguished from

other species, i.a., by the characteristically shaped posterior telomeral lamella (Fig. 29).

Etymology

The name honours Tarombera Mwabvu in recognition of his work on the taxonomy of African millipedes.

Material studied (total: 11 c?c?)

Holotype

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°06’

47.4” S, 35°56’ 50.4” E, montane, 1800-1955 m asl, plot MS - plot A, 29 Oct. 2000, Frontier Tanzania

leg. (ZMUC).

Paratypes

All from TANZANIA, Iringa Region, Udzungwa Mts: 2 SS, Iringa District, New Dabaga/Ulangambi

FR, 08°06’ 47.4” S, 35°56’ 50.4” E, montane, 1800-1855 m asl, plot MS - plot A, sample 3 of 3, 11

Nov. 2000, Frontier Tanzania leg. (ZMUC); 5 SS, Iringa District, New Dabaga/Ulangambi FR, 08°06’

47.4” S, 35°56’ 50.4” E, montane, 1800-1955 m asl, plot MS - plot A, 29 Oct. 2000, Frontier Tanzania

leg. (ZMUC); 3 Mufindi, Kigogo Forest Reserve, 1900 m, 7-15 Oct. 1984, pitfall traps in montane

rain forest, N. Scharff leg. (ZMUC).

Type locality

TANZANIA: Iringa Region, Iringa District, Udzungwa Mts, New Dabaga/Ulangambi FR, 08°06’ 47.4”

S, 35°56’ 50.4” E, montane, 1800-1955 m asl.

Description (male)

Diameter. 1.8-1.9 mm, 43—45 podous rings.

Colour. After 13 years in alcohol brownish yellow, dorsolaterally dark brown, a broad middorsal pale

stripe, a narrow blackish lateral stripe at ozopore level.

Anal valves (Fig. 6C, F). Each with a long, slightly curving dorsal spine and a small, triangular ventral

one; marginal rim raised, setiferous tubercles barely developed, not on ‘ravelins’.

Fimbus. With long, slender, apically denticulate lobes, separated by wider sinuses.

Tarsal setation. Normal.

Gonopod coxa (Figs 9,29). About 4 x as long as wide; lateral margin evenly convex in distal %; cucullus

(cu) small, smoothly rounded, without processes. Metaplical flange (inf) ending in triangular, apicad

process; metaplical mesal margin straight, but with triangular process (mp) just below level of arculus.

Metaplical shelf (ms) regularly rounded, projecting posteriad as well as mesad, continuing without

transition in metaplical shelf-spine (mss). Metaplical shelf-spine (mss) of medium length, curving first

mesad, then distad, then pointing mesad again; in specimen prepared for SEM pointing obliquely basad,

tip reaching beyond basomere.

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 29. Chaleponcus mwabvui sp. nov., right gonopod. A. Posterior view. B. Anterior-mesal view.

C. Coxa, tip, anterior view. D. Telopodite, apical-dorsal view. E. Process pip of telomere, al =

anterior telomeral lamella, cu = cucullus, mf= metaplical flange, mp = mesal metaplical process, ms =

metaplical shelf, mss = metaplical shelf-spine, pi = posterior telomeral lamella, plpl, plp2, posterior

telomeral lamella processes,/ 2 s = proximal spine on solenomere, slm = solenomere. Scales: 0.1 mm

(A-D), 0.05 mm (E).

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Gonopod telopodite (Fig. 29). Solenomere with a long, straight, mesad, proximal spine (ps). Telomere

distally with two branches:

• a large, convoluted anterior lamella ( al ) with smooth edges except for a few distal denticulations,

• a posterior-distal lamella (pi); distal margin of pi deeply laciniate in apical part, basally produced in

slender process furcating into larger branch ( plpl ) with long-spinose margins and smaller, slender

branch ( plp2 ) with large spine at base and terminally bifurcating.

Distribution and habitat

Known from two forest reserves in the Udzungwa Mts: New Dabaga/Ulangambi FR and Kigogo FR.

Altitudinal range: 1800-1955 m asl. Habitat: montane forest.

Coexisting species

All four species known from Kigogo FR, i.e., C. quasimodo sp. nov., C. scopus sp. nov., C. howelli

sp. nov. and C. mwabvui sp. nov., were found in the same samples. In New Dabaga/Ulangambi FR,

C. dabagaensis, C. gracilior sp. nov., C. netus sp. nov. and C. termini sp. nov. were found in the same

samples as C. mwabvui sp. nov. In addition, C. krai sp. nov., C. malleolus sp. nov., C. nectarinia sp. nov.,

C. teres sp. nov., C. vandenspiegeli sp. nov. and C. vilici sp. nov. occur in New Dabaga/Ulangambi FR.

Chaleponcus howelli sp. nov.

urn:lsid:zoobank.org:act:F7D97CB8-4791-4584-AD89-F2BCC6589590

Figs 4, 9, 30

Diagnosis

Medium-sized. Gonopod coxa (Fig. 9) without a lateral process; cucullus small. Distinguished from

other species, i.a., by the presence of a spine (sts) on the main stem of the telomere in combination with

the large distal spine (als) on the anterior telomeral lamella.

Etymology

The name honours Kim Howell in recognition of his huge efforts to increase the knowledge of Tanzanian

millipedes.

Material studied (total: 100 c?c?)

Holotype

TANZANIA: Iringa Region, Udzungwa Mts, Mufindi, Kigogo Forest Reserve, 1900 m, 7-15 Oct.

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

Paratypes

TANZANIA: 61 S3, Iringa Region, Udzungwa Mts, Mufindi, Kigogo Forest Reserve, 1700 m, 8-10

Oct. 1984, pitfall traps in montane rain forest, N. Scharff leg. (ZMUC); 38 S3, same data as holotype

(ZMUC).

Type locality

TANZANIA: Iringa Region, Udzungwa Mts, Mufindi, Kigogo Forest Reserve, 1900 m, montane rain

forest.

Description (male)

Diameter. 2.0-2.2 mm, 44^17 podous rings.

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 30. Chaleponcus howelli sp. nov., left gonopod. A. Mesal-ventral (-anterior) view. B. Posterior

view. C. Anterior view. D. Ventral (-anterior) view, al = anterior telomeral lamella, a Is = spine on al ,

cu = cucullus, mf= metaplical flange, mp = mesal metaplical process, ms = metaplical shelf, mss =

metaplical shelf-spine, pi = posterior telomeral lamella, ps = proximal spine on solenomere, sts = spine

on telomeral main stem. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Colour. After 30 years in alcohol almost uniform pale brown, with faint traces of a light dorsal stripe,

somet im es with a thin dark line in midline.

Anal valves. Each with a long, dorsal spine and a well-developed, but small, triangular ventral one;

marginal rim raised, setiferous tubercles well developed, on ‘ravelins’.

Fimbus (Fig. 4B). With long, slender triangular lobes, separated by rounded sinuses; lobes striate on

external surface.

Tarsal setation. Normal.

Gonopod coxa (Figs 9, 30). About 3!4 x as long as wide. Fateral margin almost straight, towards

apex gently curving mesad and continuous with apical margin; cucullus (cu) an almost perfect hollow

hemisphere. Metaplical flange (mf) ending in blunt, equilateral triangular process; metaplical mesal

margin with large rounded process (mp) delimiting semicircular sinus together with base of metaplical

shelf. Metaplical shelf (ms) simple, regularly rounded, mostly projecting laterad. Metaplical shelf-spine

(mss) of medium length, originating from anterior-mesal part of ms , projecting disto-mesad, then basad,

reaching beyond telopodital basomere.

Gonopod telopodite (Fig. 30). Solenomere with a very long, straight, meso-basad, proximal spine (ps).

Main stem of telomere with a small spine (sts) ca. in middle of basal surface. Telomere distally with two

branches:

• a rounded anterior lamella (al) ending in a long, gently sinuous mesad spine (als),

• a large, irregular posterior lamella (pi).

Distribution and habitat

Known only from Udzungwa Mts, Kigogo FR. Altitudinal range: 1700-1900 m. Habitat: montane rain

forest.

Coexisting species

All four species known from Kigogo FR, i.e., C. quasimodo sp. nov., C. scopus sp. nov., C. howelli sp.

nov. and C. mwabvui sp. nov., were found in the same samples.

Chaleponcus tintin sp. nov.

urn:lsid:zoobank.org:act:E350A221-3166-45FA-B897-F5A36A31A900

Figs 9,31,34

Diagnosis

Small. Distinguished from all other species in the C. dabagaensis group by the hook-like tip of the

gonopod coxa (Fig. 9). Gonopod coxa without a lateral process; metaplical shelf-spine long.

Etymology

The name is a noun in apposition and refers to a cartoon character developed by the Belgian author

Herge because of the (somewhat remote) resemblance of the shape of the gonopod cucullus to Tintin’s

hairstyle.

Material studied (total: 1 <$)

Holotype

TANZANIA: Iringa District, Udzungwa Mts, Chervemba, Mufu forest, Ndundulu forest, Udekwa

village, 2100 m, montane evergreen forest, in dead wood, 6 Sep. 1991, Fouis Hansen leg. (ZMUC).

European Journal of Taxonomy 100: 1-75 (2014)

Fig. 31. Chaleponcus tintin sp. nov., left gonopod. A. Anterior view. B. Mesal-anterior view. C. Ventral

(-anterior) view. D. Posterior (-mesal) view, al = anterior telomeral lamella, cu = cucullus, mfp =

metaplical flange process, ms = metaplical shelf, mss = metaplical shelf-spine, pi = posterior telomeral

lamella, plb = basal branch of pi, ps = proximal spine on solenomere. Scales 0.1 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Type locality

TANZANIA: Iringa District, Udzungwa Mts, Chervemba, Mufti forest, Ndundulu forest, Udekwa

village, 2100 m, montane evergreen forest.

Description (male)

Diameter. 1.6 mm, 41 podous rings.

Colour. After 23 years in alcohol almost uniform pale brownish, with traces of a pale dorsal band.

Anal valves. Each with a strong dorsal spine and a very small ventral one; marginal rim raised, setiferous

tubercles inconspicuous, not on ‘ravelins’.

Fimbus. With triangular lobes; lobes slightly longer than wide, striate on outer surface.

Tarsal setation. Normal

Gonopod coxa (Figs 9, 31). About 3 x as high as wide. Fateral margin straight, apically curving mesad

at right angles, cucullus (cu) produced in characteristic laterad-curving hook. Metaplical flange ending

in triangular process (mfp). Metaplical shelf (ms) small and simple; metaplical shelf-spine {mss) short,

thin, sinuous, pointing mesad.

Gonopod telopodite (Fig. 31). Solenomere with a very small proximal spine ( ps ). Telomere with two

branches:

• a relatively small, subrectangular anterior lamella (al),

• a more elaborate posterior lamella {pi) with strongly spiny margins and with a basal branch (plb)

ending in long spine; pi including plb curving anteriad and best visible in anterior view.

Distribution and habitat

The unique holotype was found in Udzungwa Mts, Chervemba, Mufii forest, Ndundulu forest, Udekwa

village. Altitude: 2100 m asl (the highest accurate record of the C. dabagaensis group). Habitat: montane

evergreen forest, in dead wood.

Coexisting species

No further species of the C. dabagaensis group is known from Ndundulu forest. However, C. basiliscus

sp. nov., C. circumvallatus sp. nov., C. gracilior sp. nov., C. ibis sp. nov. and C. netus sp. nov. also occur

in West Kilombero FR to which Ndundulu forest belongs.

Key to species of the Chaleponcus dabagaensis group

The 21 known species of the C. dabagaensis group are almost all quite distinctive in gonopod

morphology. A key to the species is given below, but comparison with the gonopod coxa profiles in Fig.

9 will often lead directly to the right species. As new species of the group are likely to be discovered,

and as the species in the key may prove to be more variable with regard to number of podous rings

and body diameter, any identification should be checked against the detailed gonopod description and

illustrations, and the size (no. of podous rings vs. body diameter) should be compared with the diagrams.

Figs. 2-3. The limbus (Fig. 4) offers additional diagnostic characters.

The key is based on adult males.

1. Body diameter >2.5 mm .2

- Body diameter <2.5 mm .5

European Journal of Taxonomy 100: 1-75 (2014)

2. Gonopod coxa with two lateral processes. basiliscus sp. nov.

- Gonopod coxa without lateral processes .3

3. Anterior lamella of telomere divided into a horizontal lobe with deeply laciniate edges and a long,

straight, distal spine-like part (Fig. 19C). Setation of walking leg tarsi normal (as Fig. 5A) ...Crafsp.nov.

- Anterior lamella of telomere different. Setation of walking legs unusual: claws surrounded by

dense ‘palisade’ of setae (Fig. 5B-C) .4

4. Posterior lamella of telomere ending in two long spines. Anterior lamella of telomere with small

accessory lamella, otherwise with smooth edges (Fig. 2 IB) . circumvallatus sp. nov.

- Posterior lamella of telomere ending in two short processes. Anterior lamella of telomere with a

coarsely dentate basal part (Fig. 22C). ibis sp. nov.

5 45—49 podous rings. Body diameter 1.5-1.8 mm (Fig. 2). Cucullus of gonopod coxa subrectangular

(Fig. 28). Limbus with spatulate lobes (Fig. 4F) . gracilior sp. nov.

- < 45 body rings and/or body diameter >1.9 mm. Cucullus and limbus different.6

6 Gonopod coxa with a lateral process .7

- Gonopod coxa without a lateral process .14

7 < 40 body rings. Body diameter 1.7-1.8 mm. No dorsal spine on anal valves. Lateral process of

gonopod coxa a strongly bent hook (Fig. 14) . malleolus sp. nov.

- > 40 body rings. Body diameter >1.8 mm. Each anal valve with a dorsal spine. Lateral process

of gonopod coxa different .8

8 Metaplical shelf of gonopod coxa in two levels between which metaplical shelf-spine fits. Lateral

process of gonopod coxa small to inconspicuous (Fig. 24) . vilici sp. nov.

- Metaplical shelf in one level. Metaplical shelf-spine and lateral process different .9

9 Lateral margin of gonopod coxa basal to lateral process pronouncedly ‘hunch-backed’ (Figs 12-13)

.10

- Lateral margin of gonopod coxa basal to lateral process not pronouncedly ‘hunch-backed’.11

10 Main stem of telomere with thumblike process (Fig. 12D) . dabagaensis

- Main stem of telomere without thumblike process (Fig. 13). quasimodo sp. nov.

11 Proximal lobe of telomere with long, fluted spine (Figs 10A, 1 IE). netus sp. nov.

- Proximal lobe of telomere without a spine.12

12 Lateral process of gonopod coxa small. Cucullus without a large mesal extension (Fig. 17)

. mwanihanensis sp. nov.

Lateral process of gonopod coxa larger. Cucullus with a large mesal extension (Figs 15-16) ... 13

13 Lateral process of gonopod coxa pointed (Figs. 15A) . scopus sp. nov.

Lateral process of gonopod coxa broadly rounded, irregularly dentate (Fig. 16B)

. nikolajscharffi sp. nov.

14 Metaplical shelf-spine of gonopod coxa long, projecting ± mesad (Figs 20A-B, 23 A, D).15

- Metaplical shelf-spine not long, projecting ± mesad.16

15 Posterior lamella of telomere divided into a horizontal lobe with spinose edges and a long,

straight, distal spine-like part (Fig. 20C, E) . nectarinia sp. nov.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Posterior lamella of telomere with strongly laciniate edges but without a long, straight, spine-like

part . vandenspiegeli sp. nov.

16 Metaplical shelf of gonopod coxa in two levels between which metaplical shelf-spine fits.

Gonopod coxa with a small to inconspicuous lateral process (Fig. 24) . vilici sp. nov.

- Metaplical shelf in one level. Metaplical shelf-spine different.17

17 Gonopod coxa with distal hook-like extension (Fig. 31A). 41 podous rings, body diameter 1.6 mm

. tintin sp. nov.

Gonopod coxa without distal hook-like extension. Mostly >41 podous rings and/or body

diameter >1.6 mm .18

18 Anal valves without dorsal spines (Fig. 6D) . teres sp. nov.

Each anal valve with a dorsal spine .19

19 Cucullus of gonopod coxa subrectangular in outline (Fig. 26B-C) . hamerae sp. nov.

- Outline of cucullus more rounded .20

20 Anterior lamella of gonopod telomere with long, smooth spine (Fig. 30C-D). howelli sp. nov.

- No such spine .21

21 Anterior lamella of gonopod telomere divided into two triangular lobes with serrate margins

(Fig. 27B, E). Posterior lamella simple . termini sp. nov.

Anterior lamella of gonopod telomere simple. Posterior lamella with a slender, spinose branch with

a long, thin, bifurcate side branch (Fig. 29E) . mwabvui sp. nov.

Distribution and habitat

Fig. 32 shows the distribution of the 21 species of the Chaleponcus dabagaensis group plotted on

a semi-diagrammatic map of the Forest Reserves (FR) from which species of the group have been

collected. Fifteen of the species have been collected in only one FR, with New Dabaga/Ulangambi FR

being the richest (6 exclusive species, 11 species in all).

Five species are shared between two FR, and C. gracilior sp. nov. even occurs in four FR, being clearly

the most widespread of the species. The easternmost FR, Mwanihana, does not share any species with

any other FR.

As Fig. 1 clearly shows, there are several forested areas in the Udzungwa Mts where the C. dabagaensis

group has not been collected. This is probably due to a collecting bias, since the very comprehensive

millipede material from the Udzungwa Mts kept in ZMUC largely derives from the FRs where the C.

dabagaensis group is now known to occur. Additional collection effort, especially in the large southern-

central Matundu forest, is likely to reveal several additional species of the group. Even renewed

collecting in the forest reserves already sampled is likely to reveal new species, since three (ibis sp.

nov., vandenspiegeli sp. nov., tintin sp. nov.) out of the 21 species are singletons and one (< circumvallatus

sp. nov.) is a doubleton (Fig. 33).

The Chaleponcus dabagaensis group consists of high-altitude species, all material having been collected

at 1390-2100 m asl, and with the highest species diversity above 1700 m (Table 2). The vast majority of

specimens was collected in montane forest, but some species were found in disturbed habitats as well,

e.g., the relatively widespread C. gracilior sp. nov. Very often several (up to 5) species were represented

in a sample, but whether such coexisting species occupy different microhabitats is not known.

European Journal of Taxonomy 100: 1-75 (2014)

Two notes on general morphology

Torsion of the gonopod telopodite

Torsion of the gonopod telopodite is characteristic of the majority of species of Odontopygidae and

the related family Spirostreptidae (Kraus 1966, Hoffman 2008) and is important for the function of the

gonopod (Barnett & Telford 1996). In all odontopygids examined by me the torsion always begins with

an anteriad bend, i.e., the torsion is clockwise when followed from coxa towards the gonopod tip and

viewed from a mesal point of view on the RIGHT gonopod. Published drawings by, e.g., Kraus (1966)

and Frederiksen (2013a,b) confirm the generality of this pattern.

Intercalary cuticular microscutes

Very small (ca. 1 pm) circular microstructures were frequently observed between cuticular microscutes

on body rings (Fig. 34). Similar structures were reported by Akkari & Enghoff (2011) from several

Fig. 32. Distribution of species of the Chaleponcus dabagaensis group in Forest Reserves of the

Udzungwa Mts. The position of the Forest Reserves in relation to each other approximately corresponds

to their true relative positions, cf. Fig. 1. Underlined species acronyms denote species known only

from one Forest Reserve. Shared species are shown on lines connecting the circles. - Forest Reserve

acronyms: KIG = Kigogo, KK = Kiranzi/Kitunguli, KR = Kisinga/Rugaro, MWA = Mwanihana, NDU =

New Dabaga/Ulangambi, US = Udzungwa Scarp, WKS = West Kilombero Scarp. - Species acronyms:

BAS = basiliscus, CIR = circumvallatus, DAB = dabagaensis , GRA = gracilior, HAM = hamerae,

HOW = howelli, IBI = ibis, KRA = krai, MAL = malleolus, MWB = mwabvui, MWN = mwanihanensis,

NEC = nectarinia, NET = netiis, NIK = nikolajscharjfi, QUA = quasimodo, SCO = scopus, TES = teres,

TEM = termini, TIN = tintin, VAN = vandenspiegeli , VIL = vilici.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

Table 2. Altitudinal range of species of the Chaleponcus dabagaensis group, based on “maximal ranges”,

see Material and methods. No altitude data are available for C. basiliscus and C. ibis.

Altitude (m)

1390-

1499

1500-

1599

1600-

1699

1700-

1799

1800-

1899

1900-

1999

2000-

2099

21 GO-

2199

netus

dabagaensis

quasimodo

malleolus

scopus

nikolajscharffi

mwanihanensis

krai

nectarinia

circumvallatus

vandenspiegeli

vilici

teres

hamerae

termini

gracilior

mwabvni

howelli

tintin

120

100

c n

| 80

v) 60

H—

f 40

U In i

tW/W W/ / 4

</// # <-* </ /

Fig. 33. The number of studied specimens (males) per species.

European Journal of Taxonomy 100: 1-75 (2014)

families of the order Polydesmida and have also been observed on Lusitanipus alternans (Verhoeff,

1893) (Callipodida: Dorypetalidae) (HE and S. Reboleira unpublished), but have not been recorded

from other millipede orders. Such intercalary cuticular microscutes are, however, discernible on fig. 8

of Frederiksen (2013a) ( Chaleponcus parensis Frederiksen, 2013), as well as on fig. 1 of Frederiksen

(2013b) (Lamelloramus rhombiformis Frederiksen, 2013b). They have also been observed on a species

of the genus Aquattuor Frederiksen, 2013, and may be widespread in Odontopygidae. The nature of the

microscutes is unknown. In a preparation of C. nikolajscharffi sp. nov. some of the microscutes give

Fig. 34. Intercalary cuticular microscutes in Chaleponcus spp. A. C. netus sp. nov. B. C. tintin sp. nov.

C-D. C. nikolaj scharffi sp. nov. Fig. D shows fungus-like outgrowths from scutes. Scales 0.001 mm.

ENGHOFF FL, Chaleponcus millipedes from Udzungwa Mountains

rise to projecting, club-shaped outgrowths of a few pm length (Fig. 34D) which may represent parasitic

fungi of a kind?

Discussion

The high diversity of odontopygids, in casu the genus Chaleponcus , in forests at high elevations in the

Udzungwa Mountains is impressive and somewhat unexpected: odotopygids in general “do not seem to

be especially abundant at higher elevations” (Hoffman 1993). Collections of odontopygids at hand from

other Eastern Arc mountain blocks are much less diverse.

All 21 Udzungwan Chaleponcus species belong to the morphologically well-defined C. dabagaensis

group which is most probably monophyletic as suggested by the unique, potentially autapomorphic

spine-bearing shelf on the gonopod coxa. No species referable to this group is known from elsewhere,

and the C. dabagaensis group can therefore be regarded as an endemic high altitude Udzungwan ‘species

swarm’. To my knowledge, there are no similar examples of taxa which have radiated ‘sympatrically’

within the Udzungwa Mountains.

Too little microhabitat information is available to support a hypothesis of adaptive radiation similar to

that observed in certain insular species swarms of millipedes (Enghoff 1983, 2011b; Enghoff & Baez

1993). However, the frequent occurrence of multiple species in a sample (Table 1), in combination with

the considerable inter-specific differences in body size (Figs 2-3), as well as the unusual tarsal setation

of a few species (C. circumvallatus sp. nov., C. ibis sp. nov.), indicates that adaptive radiation may

indeed have taken place.

Acknowledgements

This work would not have been possible at all without the impressive collections made by Frontier

Tanzania. My sincere thanks are due to this NGO, as well as to Nikolaj Scharff and other colleagues at

the Natural History Museum of Denmark who have also collected important material and have provided

various advices. Further thanks are due to Judith Marshall and her colleagues in the Virginia Museum

of Natural History for taking good care of me during my visit and for arranging loan of specimens from

the collection of the late Richard Hoffman, to Fuca Bartolozzi for loaning specimens from the Firenze

museum, to Andy Marshall for advice, and to Nesrine Akkari and Sergei Golovatch for very useful

co mm ents on the manuscript.

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Manuscript received: 11 August 2014

Manuscript accepted: 24 September 2014

Published on: 24 October 2014

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

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, De nm ark