European Journal of Taxonomy 177: 1-19

http://dx.doi.org/10.5852/ejt.2016.177

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

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Enghoff H.

Research article

urn:lsid:zoobank.org:pub:84816D6D-B8EE-4D90-A6DD-77C86EAC7CDB

A mountain of millipedes III: A new genus for three new species from

the Udzungwa Mountains and surroundings, Tanzania, as well as

several ‘orphaned’ species previously assigned to Odontopyge

Brandt, 1841 (Diplopoda, Spirostreptida, Odontopygidae)

Henrik ENGHOFF

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

Kobenhavn 0, Denmark, email: [email protected]. dk

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

Abstract. The new genus Geotypodon gen. nov. is described. It includes two species from the Udzungwa

Mountains: G. millemanus gen. et sp. nov. (type species) and G. submontanus gen. et sp. nov., one

species from nearby Iringa: G. iringensis gen. et sp. nov., and 18 previously described species hitherto

incorrectly assigned to Odontopyge Brandt, 1841.

Keywords. Eastern Arc, taxonomy, new species.

Enghoff H. 2016. A mountain of millipedes III: A new genus for three new species from the Udzungwa Mountains

and surroundings, Tanzania, as well as several ‘orphaned’ species previously assigned to Odontopyge Brandt, 1841

(Diplopoda, Spirostreptida, Odontopygidae). European Journal of Taxonomy 177: 1-19. http://dx.doi.org/10.5852/

eit.2016.177

Introduction

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

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

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

Frederiksen (2015).

In the present article three further new odontopygid species are described from the Udzungwa Mountains

and their immediate surroundings (Fig. 1). The new species resemble several species which were

assigned to the genus Odontopyge Brandt, 1841 by Kraus (1960) and authors before him. However,

Hoffman (1991) showed that Odontopyge had been misinterpreted by virtually everybody dealing with

the genus after its original description and that Odontopyge is the correct name for a genus until then

known as Omopyge Manfredi, 1941. Hoffman (1991) rectified the situation and formally synonymized

Omopyge under Odontopyge , but by this action he left a large number of species originally described in

or subsequently transferred to Odontopyge ‘orphaned’, i.e., without a valid genus name. Some of these

have been transferred to other genera by Hoffman & Howell (1981, 2012), Hoffman (2000, 2002) and

Frederiksen & Enghoff (2015), but very many remain orphaned.

European Journal of Taxonomy 177: 1-19 (2016)

Kraus (1960) was aware of the situation and therefore accepted Attems ’ (1909) designation of Odontopyge

kilimandjaronus Attems, 1909, as type species of Odontopyge , mentioning that the ICZN would be

asked to make Attems’ designation valid. This, however, has not happened, and Attems’ type designation

remains invalid, as does that of Silvestri (1896: Jidus bicuspidatus Brandt, 1841) (Jeekel 1970).

Table 1 lists all species assigned to Odontopyge by Kraus (1960) and subsequent authors, with an

indication of their subsequent re-classification. Part 2 of the table includes the dubious species of

6 Odontopyge ’ listed by Kraus (1960) - many of these names were based on female specimens and may

never be resolvable.

To accommodate the new species described here, a new genus is established to which several of the

orphaned species can be assigned as well.

Material and methods

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

Denmark, University of Copenhagen (ZMUC). The specimens were partly collected during field trips

TANZANIA

MIKUMI

IRINGA

W Kilombero

r Scarp

Nyambanito

Ukami

Kiranzi /

Kitungul

igulu

Wooded grassland

Woodland

| Open forest

|9 Closed forest

O Towns

Major roads

Rivers

Fig. 1 . Map of the Udzungwa Mountains, showing the collecting sites for the three new Geotypodon

species, as well as names of the Forest Reserves in question and names of individual mountains in

West Kilombero FR. Red diamonds = G. millemanus gen. et sp. nov., yellow dot: G. submontanus gen.

et sp. nov., blue triangle: G. iringensis gen. et sp. nov. Based on fig. 1 in Marshall et al. (2010) and

information in Doody et al. (2001).

ENGHOFF H., New millipede genus and species from the Udzungwa Mts

Table 1 . Species of Odontopyge sensu auctorum nee Brandt, 1841. The table is based on Kraus (1960),

with updates as indicated. Part 1 of the list contains species placed by Kraus and subsequent authors in

Odontopyge. Part 2 contains the “ nomina dnbicT listed by Kraus (1960). Numerous additional species

originally described in Odontopyge and transferred to other genera by Kraus and his predecessors are

not listed.

Part 1. Species assigned to ‘ Odontopyge sensu auctorum’ by Kraus and subsequent authors.

Odontopyge

Species

Current placement Reference

angolana Kraus, 1958

antrophila Attems, 1951

arrogans (Attems, 1935)

astragalus (Attems, 1912)

bayoni Silvestri, 1910

carli Kraus, 1960

citernii S ilvestri, 1910

delitescens Attems, 1935

dewittei Kraus, 1958

difficilis Silvestri, 1895

dispersa Carl, 1909

dorsalis Carl, 1909

errata Kraus, 1960

franco Sudani c a Attems, 1914

gracilitarsus Kraus, 1958

heteromodesta Kraus, 1960

indecisus Pierrard, 1970

intermedia Carl, 1909

kakandae Kraus, 1958

kandti kandti Carl, 1909

kandti denticulata Attems, 1937

kilembeensis Demange, 1990

kilimanjarona Attems, 1909

latifolia (Attems, 1914)

leviceps (Attems, 1909)

medjensis (Chamberlin, 1927)

meneliki Attems, 1927

multianulata Attems, 1914

ollieri Silvestri, 1907

paludosai Attems, 1953)

par dal is (Gerstacker, 1873)

picea Attems, 1938

procera Attems, 1914

procerula Kraus, 1960

pnnctulata Attems, 1912

scaphula Attems, 1912

sennae Brolemann, 1903

severini Silvestri, 1897

simplex (Chamberlin, 1927)

Geotypodon comb. nov.

‘orphaned’

Geotypodon comb. nov.

‘orphaned’

Geotypodon comb. nov.

Calyptomastix

Geotypodon comb. nov.

‘orphaned’

Geotypodon comb. nov.

Calyptomastix

‘orphaned’

Rhamphidarpoides

Callistodontopyge

Calyptomastix

‘orphaned’

Geotypodon comb. nov.

‘orphaned’

Calyptomastix

Geotypodon comb. nov.

‘orphaned’

Geotypodon comb. nov.

‘orphaned’

Hoffman (2012)

Pierrard (1970)

Hoffman (2012)

Demange (1990)

Frederiksen & Enghoff (2015)

Hoffman & Howell (1981)

Hoffman (2012)

European Journal of Taxonomy 177: 1-19 (2016)

Odontopyge

specularis Attems, 1927

Geotypodon comb. nov.

Odontopyge

stenotarsa Attems, 1938

Geotypodon comb. nov.

Odontopyge

terebrum Ribaut, 1907

‘orphaned’

Odontopyge

turn idem Karsch, 1881

‘orphaned’

Odontopyge

uv ire ns is Kraus, 1960

‘orphaned’

Odontopyge (?) vanutellii Silvestri, 1898

‘orphaned’

Part 2. Nomina dubia listed by Kraus (1960).

Original genus

(subgenus) _

Spirostreptus

(Odontopyge)

Spirostreptus

(Odontopyge)

Odontopyge

Ildus (Spirostreptusj

Spirostreptus

Ctenoiulus

Odontopyge

lulus (Spirostreptus)

Spirostreptus

(Odontopyge)

Spirostreptus

(Odontopyge)

Odontopyge

lulus (Spirostreptus)

Odontopyge

lulus (Spirostreptus)

Odontopyge

Species Current placement Reference

acutus Karsch, 1881

aequalis Porath, 1982

amaura Brolemann, 1926

angolensis Karsch, 1881

anomala Silvestri, 1896

attenuata Silvestri, 1895

bicolor Silvestri, 1897

bicuspidatus Brandt, 1841

binodiferNogQS, 1878

chatelainei Cook, 1893 Ctenoiulus Hoffman (1980)

dilatata Brolemann, 1905

dimidiatiformis Porat, 1893

diversicolor Silvestri, 1895

diversifacies Silvestri, 1898

doriae Silvestri, 1896

ecarinata Porat, 1894

exquisita Silvestri, 1896

flavotaeniatus Brandt, 1841

foveolatus Porath, 1872

r T7- i i ooi Ctenoiulus chatelainei TT .. AOA n

furcatus Karsch, 1881 „ . Honman (1980)

Cook, 1893

gestrii Silvestri, 1896

gracilicornis Brandt, 1841

jallae Silvestri, 1896

hollarii Brandt, 1841 Odontopyge Hoffman (1991)

litoranea Silvestri, 1897

ENGHOFF FL, New millipede genus and species from the Udzungwa Mts

Spirostreptus

(Odontopyge)

Spirostreptus

(Odontopyge)

Spirostreptus

(Odontopyge)

Odontopyge

Spirostreptus

(Odontopyge)

Odontopyge

maculatus Karsch, 1881 Callistodontopyge

m itellatus Karsch, 1881 Callistodontopyge

praetextus Porath, 1872

puerilla Daday, 1889

puncticaudisVomXh , 1872

rubripes Silvestri, 1895

Hoffman & Howell

(1981)

Hoffman (2002)

Odontopyge

ruspolii Silvestri, 1896

Spirostreptus

Odontopyge

scaliger Gerstacker, 1873

trivial is var. strigulosa Porat,

1891

Odontopyge

Spirostreptus

Odontopyge

subelegans Silvestri, 1897

sugillatus Gerstacker, 1873 Callistodontopyge

trivial is Porat, 1894

Hoffman & Howell

(1981)

Odontopyge

uebicola Silvestri, 1896

by ZMUC staff and students, partly by the NGO Frontier Tanzania (see Marshall et al. 2001). A total of

seven male specimens was examined. All are kept in 70% alcohol.

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

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

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

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

See Enghoff (2014) for the description standards used.

Abbreviations for morphological terms used in the descriptions and on illustrations

atl = anterior distal lobe of telomere

bl = basal lamella of telomere

cx = coxa

itl = intermediate distal lamella of telomere

Ic = lateral concavity of coxa

Ifl = longitudinally folded lamella

// = longitudinal lamella

mf = anteriad metaplical flange

mla = metaplical lamella

mp = metaplica

msp = metaplical spine-like process

pn = posttorsal narrowing

pp = proplica

ptl = posterior distal lobe of telomere

pts = proximal telomeral spine

European Journal of Taxonomy 177: 1-19 (2016)

slm = solenomere

tl = terminal lobe of telomere

tt = torsotope

Abbreviations used in the text, other than the above

asl = above sea level

FR = Forest Reserve

ZMUC = Natural History Museum of Denmark (Zoological Museum)

Results

Taxonomy

Class Diplopoda Blainville-Gervais, 1844

Order Spirostreptida Brandt, 1833

Family Odontopygidae Attems, 1909

Subfamily Archepyginae Manfredi, 1939

Tribe Prionopetalini Hoffman, 1991

Geotypodon gen. nov.

um:lsid:zoobank.org:act:54706BDE-AF06-42BC-A2B3-F2D23A097B28

Type species

G. millemanus gen. et sp. nov.

Diagnosis

A genus of Odontopygidae-Prionopetalini characterized by: a long basad metaplical spine on the anterior

side of the coxa, a compact torsotope, lack of pretorsal or torsal spines/processes, a pronounced posttorsal

narrowing without spines, a division of the telopodite into solenomere and telomere immediately distal

to posttorsal narrowing, a spine emerging from the base of the telomere and curving more or less parallel

to the basal parts of the solenomere, a slender, whip-like, smooth solenomere without any outgrowths or

appendages (except sometimes a tiny subapical spine), a highly three-dimensional telomere consisting

of various lobes and lamellae with largely smooth margins.

Etymology

An anagram of Odontopyge. To be regarded as masculine in analogy with other names ending in -odon.

Other species included:

G. angolanus (Kraus, 1958) (comb. nov. ex Odontopyge )

G. bayoni (Silvestri, 1910) (comb. nov. ex Odontopyge)

G. carli (Kraus, 1960) (comb. nov. ex Odontopyge )

G. dispersus (Carl, 1909) (comb. nov. ex Odontopyge)

G. erratus (Kraus, 1960) (comb. nov. ex Odontopyge )

G. gracilitarsus (Kraus, 1958) (comb. nov. ex Odontopyge)

G. heteromodestus (Kraus, 1960) (comb. nov. ex Odontopyge)

G. indecisus (Pierrard, 1970) (comb. nov. ex Odontopyge)

G. intermedius (Carl, 1909) (comb. nov. ex Odontopyge)

G. iringensis gen. et sp. nov.

G. multianulatus (Attems, 1914) (comb. nov. ex Odontopyge)

G. ollieri (Silvestri, 1907) (comb. nov. ex Odontopyge)

ENGHOFF FL, New millipede genus and species from the Udzungwa Mts

G. piceus (Attems, 1938) (comb. nov. ex Odontopyge)

G. procerus (Attems, 1914) (comb. nov. ex Odontopyge)

G. procerulus (Kraus, 1960) (comb. nov. ex Odontopyge )

G. punctulatus (Attems, 1912) (comb. nov. ex Odontopyge)

G. sennae (Brolemann, 1903) (comb. nov. ex Odontopyge)

G. specular is (Attems, 1927) (comb. nov. ex Odontopyge)

G. stenotarsus (Attems, 1938) (comb. nov. ex Odontopyge)

G. submontanus gen. et sp. nov.

Descriptive notes

Species of Geotypodon gen. nov. have a typical odontopygid habitus (Fig. 2) and are mostly medium¬

sized by odontopygid standards (cf. Kraus 1966). Published numbers of podous rings range from 49

(G. ollieri) to 72 (G. multianulatus) (one has been subtracted from the published numbers because

these include the telson), and published male body diameters range from 1.7 mm (G. sennae) to 5.5 mm

(G. specularis). The majority of species have 54-66 podous rings and a diameter of 4.3-5.5 mm, but

there is a group of smaller species (G. angolanus , G. erratus , G. heteromodestus , G. ollieri) with 49-54

podous rings and a diameter of 2.2-3.2 mm. G. sennae is a particularly slender species: 66-68 podous

rings and 1.7 mm diameter, and G. multianulatus , true to its name (at least as misspelled by Kraus

(1960): multiannulatus), stands out with 72 podous rings and 4.8 mm diameter. Of the new species

described here, G. millemanus gen. et sp. nov. falls neatly in the main group, G. submontanus gen. et

sp. nov. is slightly more slender than the main group, whereas G. iringensis gen. et sp. nov. has a larger

body diameter than any other described congener (Fig. 3).

Several of the species assigned to Geotypodon gen. nov. have the same type of limbus (with hand¬

like lobes) as the type species; this is true of G. angolanus , G. carli , G. erratus , G. gracilitarsus,

G. procerulus and G. punctulatus (Attems 1912; Kraus 1958, 1960) as well as G. submontanus gen. et

sp. nov. Other species have other types of limbus: G. heteromodestus , G. indecisus, G. multianulatus, G.

piceus, G. sennae, G. specularis and G. stenotarsus (Attems 1914, 1927, 1938, 1953 [G. heteromodestus,

as Haplothysanus modestus]; Brolemann 1903; Pierrard, 1970), as well as G. iringensis gen. et sp. nov.

The limbus of procerus somewhat but not quite resembles that of G. millemanus gen. et sp. nov. (Attems

1914), and no information is available about this character in G. bayoni, G. dispersus or G. intermedius.

On the other hand, several species outside Geotypodon gen. nov. as here defined have the same type

of limbus as G. millemanus gen. et sp. nov.. This is, for example, the case for several species of

Rhamphidarpoides Kraus, 1960, including R. kilimandjarona (Attems, 1909), R. ruandensis Kraus,

1960, and R. regina (Carl, 1909) (Attems 1914; Kraus 1960; Frederiksen & Enghoff 2015), Helicochetus

spp. (e.g., Kraus 1966: figs 83-87), Solenozophyllum kazibaense Kraus, 1958 and 6 Odontopyge" dewittei

Kraus, 1958.

Remarks

Although Geotypodon gen. nov. is proposed to accommodate several species of Odontopyge sensu Kraus

(1960), its diagnosis and circumscription are narrower. Species of Odontopyge sensu Kraus (1960), may

thus lack a long basad metaplical spine on the anterior side of the coxa (present in Geoptypodon), may

have spines in the torsal region (absent in Geotypodon gen. nov.), and may lack a spine emerging from

the base of the telomere and curving more or less parallel to the basal parts of the solenomere (present

in Geotypodon gen. nov.).

None of the characters listed in the diagnosis are exclusive to Geotypodon gen. nov.: Notably, the

conspicuous coxal metaplical anterior spine is also found in, e.g., several species of Rhamphidarpoides

and Raduliverpa Frederiksen & Enghoff, 2015 (Frederiksen & Enghoff 2015) as well as Spinotarsus

Attems, 1909 (Kraus 1960, 1966), but the species in question differ from Geotypodon gen. nov. by

European Journal of Taxonomy 177 : 1-19 ( 2016 )

either having the solenomere with spines or fluting ( Rhamphidarpoides , Raduliverpa ) or by having

characteristic structures on the telomere (“Basallamelle” and/or “Bogenlamelle” sensu Kraus 1960)

( Spinotarsus ).

A spine emerging from the base of the telomere is also found in several other ’ Odontopyge’ species,

e.g., citernii Silvestri, 1898, difficilis Silvestri, 1895, and severini , Silvestri, 1897 - see Kraus (1960),

where this spine is called “Tibialdorn”. It is also found in Rhamphidarpoides species (Frederiksen &

Enghoff 2015), in Calyptomastix kakandae (Kraus, 1958), as well as several Spinotarsus species, etc.

In the Chaleponcus dab ague ns is-group there is a spine at almost the same place, but emerging from the

base of the solenomere instead of the base of the telomere (Enghoff 2014).

Considering the notorious mosaic-like distribution of morphological characters throughout the family,

the genus Geotypodon gen. nov. as defined here is quite possibly not a monophyletic group, cf. the

Discussion section (see below). Until a more satisfactory analysis of relationships within Odontopygidae

becomes available, the new genus can, however, serve as a ‘home’ for several (but not all) ‘orphaned’

species hitherto classified in Odontopyge.

Geotypodon millemanus gen. et sp. nov.

urn:lsid:zoobank.org:act:D65889F9-1063-4FBB-AEBB-A594052220Bl

Figs 1-4

Diagnosis

A species of Geotypodon gen. nov. in which the gonopod coxal metaplica is produced mesad in a thin

lamella, a small longitudinal lamella, perpendicular to the other, is present on the anterior part of the

Photo: Nikolas loannou, feb 2009

Fig. 2. Geotypodon millemanus gen. et sp. nov., paratype from West Kilombero Scarp FR after nine

years in alcohol. Photograph by N. loannou.

ENGHOFF H., New millipede genus and species from the Udzungwa Mts

metaplica, the telomere is apically divided into two lobes of approximately equal size, and the limbus

lobes are multi-cusped, hand-like. It shares these characters with G. submontanus gen. et sp. nov., but

differs from that species in larger size, straight mesal margin of metaplical lamella and absence of a

spine-like process on the posterior apical telomeral lobe.

Etymology

The name is a Fatin noun in apposition, meaning “a thousand hands” and referring to the hand-like

limbus lobes. A specimen with a diameter of 5 mm will have a body perimeter of approximately 15 mm

Assuming that the limbus covers only 10 mm, and taking into account that each “hand” occupies about

10 microns of the perimeter, each body ring will carry about a thousand “hands”, and a specimen with

60 body rings will therefore have roughly 60,000 “hands”.

Material studied (total: 4 c?c?)

Holotype

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

36°22T7.6" E, montane forest, 1390-1410 m asl. Plot Paradiso, casual, 18 Nov. 2000, Frontier Tanzania

leg. (ZMUC00040350).

Paratypes

TANZANIA: 2 SS, same data as holotype (ZMUC100969, ZMUC00040345); 1 S, Iringa Region,

Iringa District, Udzungwa Mts, Kiranzi-Kitungulu FR, 08°09' S, 35°05' E, forest, 1500 m asl, Jan. 1996,

M. Andersen, P. Gravlund & A. Jakobsen leg. (ZMUC00046991).

Number of podous rings

Fig. 3. Body size of males of Geotypodon spp. Bold symbols indicate numbers of podous rings and

midbody vertical diameter of the new species described here. Small circles and shaded areas indicate

published measurements for other Geotypodon species.

European Journal of Taxonomy 177 : 1-19 ( 2016 )

Fig 4. Geotypodon millemanus gen. et sp. nov., paratype from Kiranzi-Kitungulu FR. A-E: Right

gonopod. A. Posterior view. B. Anterior view, telomere in red oval. C. (Posterior-)mesal view, solenomere

(yellow) and proximal telomeral spine (green) coloured. D. (Anterior-)mesal view. E. Telomere (part of

coxa at lower left), basal (dorsal) view. F. Limbus. Abbreviations: atl = anterior distal lobe of telomere;

bl = basal lamella of telomere; itl = intermediate distal lamella of telomere; ll = longitudinal lamella;

mf= anteriad metaplical flange; mla = metaplical lamella; mp = metaplica; msp = metaplical spine-like

process; pn = posttorsal narrowing; pp = proplica; ptl = posterior distal lobe of telomere; pts = proximal

telomeral spine; slm = solenomere; tt = torsotope. Scales: A-E = 0.1 mm, F = 0.01 mm.

ENGHOFF H., New millipede genus and species from the Udzungwa Mts

Type locality

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

36°22T7.6" E, montane forest, 1390-1410 m asl, cf. Marshall et al. (2001).

Description

Size. Fength ca. 9 cm. Diameter 4.8-5.1 mm. 60-66 podous rings, no apodous rings in front of telson.

Colour. After 15 years in alcohol somewhat faded, but pattern still evident. Upper part of head blackish,

lower part yellowish. Overall colour of body ventrally and laterally yellowish. Posterior ca. 40% of

metazona amber, in front of amber zone a blackish zone; blackish zone narrow laterally, becoming wider

dorsally where extending onto posterior V4-V3 of prozona; resulting mid-dorsal dark band becoming

narrower towards head. Telson blackish with yellowish margins, legs yellowish.

Head. Without peculiarities.

Collum. With a marginal and a submarginal furrow.

Body rings. Almost perfect cylinders, not vaulted; suture straight; ozopores ca. three diameters behind

suture.

Fimbus (Fig. 4). Consisting of isolated hand-like lobes of ca. 10 microns’ length, each with 3-6 ‘fingers’

and each with a ridge running along its length.

Anal valves. Each with a well-developed dorsal spine and a smaller, yet distinctive ventral one; margin

raised, with 3 setae not borne on tubercles.

Male legs. From 5 th pair with postfemoral and tibial pads on all legs, except last four pairs; tibial pads

absent from several pairs in front of these.

Gonopod coxa (Fig. 4). Slender, slightly sigmoid. Proplica ( pp ) ending in small proplical lobe hidden

behind anterior edge of metaplical lamella ( mla ), on Fig. 4B. Metaplica ( mp ) with poorly developed

basal anteriad flange (mf ), disto-mesally expanded into thin lamella (mla), with a straight mesal margin,

tip of metaplica formed by blunt-triangular extension of mla', a second smaller longitudinal lamella (11)

on anterior surface of metaplica partly covering proplical lobe; metaplica at level of proplical lobe, with

a long, slightly curved spine-like process (msp) directed toward base of coxa on its anterior side.

Gonopod telopodite (Fig. 4). Arculus 90°. Torsotope (tt) simple, compact, without processes (Fig. 4B).

Posttorsal narrowing (pn ) pronounced, very slender, without processes or spines (Fig. 4B). Telopodite

just distal to posttorsal narrowing dividing into slender, whip-like solenomere and complicated telomere.

Solenomere (slm) curved in 3 dimensions (at least on preserved specimens), apically pointed, without

any outgrowths (Fig. 4A, C, E). Efferent groove continuing from posttorsal narrowing onto solenomere

and running all the way to its tip. Telomere with long, stout basal spine (pts ), spine first curving in parallel

with solenomere, but then becoming straight and directed mesad (Fig. 4A, C). Telomere close to pts with

a basal lamella (bl) (Fig. 4A), followed by several complicated lamellar parts; basal part of telomere

partly sheathing base of solenomere, distal part divided into two equally-sized thin lobes (atl and ptl),

which initially diverge at ca. 90° but then curve towards each other (Fig. 4B, D-E). An intermediate

lamella (itl) lodged in the space between atl and plm (Fig. 4D). Surfaces of atl and ptl facing each other,

concave, margins of each lobe subparallel, smooth; telomere entirely without denticles or spines.

European Journal of Taxonomy 177: 1-19 (2016)

Distribution and habitat

Known from West Kilombero FR and Kiranza-Kitungulu FR. Altitudinal range: 1145-1500 m asl.

Haibtat: (montane) forest.

Coexisting species

In Kiranza-Kitungulu FR G. millemanus gen. et sp. nov. was found in the same sample as Chaleponcus

dabagaensis Kraus, 1958 and C. gracilior Enghoff, 2014. In West Kilombero FR no other odontopygids

were found in the same sample as G. millemanus gen. et sp. nov., but G. submontanus gen. et sp. nov.,

Chaleponcus basiliscus Enghoff, 2014, G. circumvallatus Enghoff, 2014, G. gracilior , G. ibis Enghoff,

2014, G. netus Enghoff, 2014, G. tintin Enghoff, 2015, Aquattuor longipala Enghoff 2015 and

A. udzungwensis Enghoff, 2015 also occur in West Kilombero FR.

Geotypodon submontanus gen. et sp. nov.

urn:lsid:zoobank.org:act:02DBFF6A-56Al-4EB3-9001-AF063F22ClB6

Figs 1, 3, 5

Diagnosis

A species of Geotypodon gen. nov. in which the gonopod coxal metaplica is produced mesad in a thin

lamella, a small longitudinal lamella, perpendicular to the other, is present on the anterior part of the

metaplica, the telomere is apically divided into two lobes of approximately equal size, and the limbus

lobes are multi-cusped, hand-like. Shares these characters with G. millemanus gen. et sp. nov., but

differs from that species in smaller size, bicuspid mesal margin of metaplical lamella and presence of a

spine-like process on the posterior apical telomeral lobe.

Etymology

The name is a Latin adjective referring to the habitat.

Material studied (total: 1 f)

Holotype

TANZANIA: <$, Iringa Region, Iringa District, UdzungwaMts, West Kilombero Scarp FR, 07°53T9.5" S,

36°23'11.6" E, submontane forest, 1145 m asl, trapsite Ukami, casual, Nov. 2000, Frontier Tanzania leg.

(ZMUC00046992).

Type locality

TANZANIA, Iringa Region, Iringa District, Udzungwa Mts, West Kilombero Scarp FR, 07°53T9.5" S,

36°23'11.6" E, submontane forest, 1145 m asl, cf. Marshall et al. (2001).

Description

Size. Length ca. 7 cm. Diameter 3.8 mm. 60 podous rings, no apodous rings in front of telson.

Other characters. As in G. millemanus gen. et sp. nov., with the following exceptions:

- telson (after 15 years in alcohol) yellowish,

- postfemoral and tibial pads on male legs smaller,

- metaplical lamella ( mla ) produced mesad, with two sharp angles (Fig. 5A-B),

- posterior distal lobe of telomere (ptl) apically with a dark, spine-like process (Fig. 5E).

Distribution and habitat

Known only from West Kilombero FR. Altitude: 1145 m asl. Habitat: submontane forest.

ENGHOFF H., New millipede genus and species from the Udzungwa Mts

Fig. 5. Geotypodon submontanus gen. et sp. nov., holotype. A-E. Feft gonopod. A. Anterior view.

B. Posterior view. C. Apical part of coxa and proximal part of telopodite, anterior view. D. (Anterior-)

mesal view. E. Posterior distal lobe of telomere; insertion highlights spine-like process. F. Fimbus.

Abbreviations: atl = anterior distal lobe of telomere; itl = intermediate distal lamella of telomere; ll =

longitudinal lamella; mla = metaplical lamella; msp = metaplical spine-like process; ptl = posterior distal

lobe of telomere; pts = proximal telomeral spine. Scales: A-D = 0.1 mm, E = 0.05 mm, F = 0.01 mm

European Journal of Taxonomy 177: 1-19 (2016)

Coexisting species

No other odontopygids were found in the same sample as G. submontanus gen. et sp. nov., but

G. millimanus gen. et sp. nov., Chaleponcus basiliscus Enghoff, 2014, C. circumvallatus Enghoff 2014,

C. gracilior , C. ibis Enghoff 2014, C. netus Enghoff, 2014, C. tintin Enghoff, 2015 , Aquattuor longipala

Enghoff, 2015 and A. udzungwensis Enghoff, 2015 also occur in West Kilombero FR.

Geotypodon iringensis gen. et sp. nov.

um:lsid:zoobank.org: act: 0171D40F - 1E0D-4194-8D1B-045D46F92095

Figs 1, 3, 6

Diagnosis

A species of Geotypodon gen. nov. in which the gonopod coxal metaplica is apically rounded, the

telomere is apically divided into a small posterior lobe and a large anterior lobe, which gives rise to a

strongly curved terminal lobe, and the limbus lobes are pointed triangular.

Etymology

The species is named after the type locality.

Material studied (total: 2 SS)

Holotype

TANZANIA: <$, IringaRegion, 10 km E of Iringa city, 7°46' S, 35°42' E, Mar.-Apr. 1996, F.F. Sorensen

leg. (ZMUC00046993).

Paratype

TANZANIA: 1 S, same data as holotype (ZMUC00046994).

Type locality

TANZANIA: Iringa Region, 10 km east of Iringa city, 7°46' S, 35°42' E.

Description

Size. Length ca. 9 cm. Diameter 6.0-6.1 mm. 61-67 podous rings, no apodous rings in front of telson.

Colour. After 19 years in alcohol uniform greyish; posterior part of metazona amber, legs and antennae

dark brownish.

Head. Without peculiarities.

Collum. With a marginal and a submarginal furrow.

Body rings. Almost perfect cylinders, not vaulted; suture straight; ozopores ca. three diameters behind

suture.

Limbus (Fig. 6). With sharply pointed lobes.

Anal valves. Each with a well-developed dorsal spine and a smaller, yet distinctive ventral one, margin

raised, with 3 setae not borne on tubercles.

Male legs. From 4 th pair with postfemoral and tibial pads on all legs, except last few pairs where only

postfemoral pads are present.

ENGHOFF FL, New millipede genus and species from the Udzungwa Mts

Fig. 6. Geotypodon iringensis gen. et sp. nov. A-D. Holotype, left gonopod. A. Anterior view. B. Posterior

view. C. Mesal-ventral view. D. Telomere and solenomere, basal (dorsal) view. — E. Paratype, limbus.

Abbreviations: atl = anterior distal lobe of telomere, bl = basal lamella of telomere, ex = coxa (seen

from the basis, with remains of muscles); itl = intermediate distal lamella of telomere; Ic = lateral

concavity of coxa; Ifl = longitudinally folded lamella; mf= anteriad metaplical flange; mp = metaplica;

msp = metaplical spine-like process; pn = posttorsal narrowing; pp = proplica; ptl = posterior distal

lobe of telomere; pts = proximal telomeral spine; slm = solenomere; tl = terminal lobe of telomere; tt =

torsotope. Scales: A-D = 0.2 mm, E = 0.01 mm

European Journal of Taxonomy 177: 1-19 (2016)

Gonopod coxa (Fig. 6). Basally parallel-sided, with antero-lateral concavity ( Ic ) (Fig. 6A). Proplica

ending in small proplical lobe (hidden behind anterior edge of metaplica on Fig. 6A). Metaplica with

poorly developed basal anteriad flange ( mf) (Fig. 6A), distally regularly rounded and projecting laterad

as semicircular lobe; metaplica at level of proplical lobe, with a long, latero-basad process ( msp ) on

anterior side of coxa (Fig. 6A-B); process straight and slender in anterior view, slightly curved and

broader in lateral view.

Gonopod telopodite (Fig. 6). Arculus 90°. Torsotope (tt) simple, compact, without processes (Fig. 6A).

Posttorsal narrowing (pn) pronounced, very slender, without processes or spines (Fig. 6A). Telopodite

just distal to posttorsal narrowing dividing into slender, whip-like solenomere and complicated telomere.

Solenomere ( slm ) curved in 3 dimensions (at least on preserved specimens), apically pointed, without

any outgrowths (Fig. 6A-D). Efferent groove continuing from posttorsal narrowing onto solenomere

and running all the way to its tip. Telomere with short, dark basal spine (pts) (Fig. 6B) and a basal

lamella ( bl) (Fig. 6A), followed by a slender part formed by a longitudinally folded lamella (If 7) with

irregular edges (Fig. 6B-C); distal part divided into a slender posterior lobe (ptl ) and a large, broad

anterior lobe (atl) (Fig. 6A-B, D), the latter terminally giving rise to a slender, strongly curved terminal

lobe (tl, curvature not visible on Fig. 6D). An intermediate lamella (itl) lodged in the space between atl

and plm (Fig. 6B).

Distribution and habitat

Known only from the environs of Iringa city. The altitude of the type locality will be at ca. 1600 m asl.

Coexisting species

No other odontopygid species were found together with G. iringensis gen. et sp. nov.

Notes

Geotypodon iringensis gen. et sp. nov. is very similar to G. multianulatus from Kenya, but there are

differences, including that in G. multianulatus the gonopod coxa has a much larger, narrower and less

rounded lateral lobe, and the solenomere has a small subdistal spine (cf. Discussion section).

Discussion

Of the three species described here, G. millemanus gen. et sp. nov. and G. submontanus gen. et sp. nov.,

both from the Udzungwa Mountains proper, are particularly similar, notably sharing the lamellar

structure of the mesal part of the coxal metaplica and the apically divided telomere with an intermediate

lamella (itl) between the apical and posterior distal lobes (atl and ptl). G. iringensis gen. et sp. nov. does

not have the metaplical lamella, but its telomere has the same distal elements (atl, itl, ptl) as the two

other species. In most other species of Geotypodon gen. nov. as here defined, the telomere does not show

a similar apical subdivision. Exceptions are G. multianulatus, G. sennae and G. specularis, where the

telomere is divided into two large lobes (Attems 1914, 1927; Brolemann 1903).

The unsatisfactory state of odontopygid taxonomy was characterized as follows by Hoffman & Howell

(2012): “That classification of odontopygid millipeds remains in a highly unsettled condition is due

both to the inherent complexity of the male genitalia and the traditional reliance on a few obvious

key-characters (a priori definition) instead of groupings made on the basis of overall similarity of the

appendages. Either approach is further complicated by frequent contradictory states of characters as

expressed in coxal or telopodital regions. Almost identical coxal forms may recur randomly amongst

taxa defined on the basis of the telopodite and thought to be not closely related. Body form tends to be

of monotonous similarity throughout the group, and female genitalia have so far provided very few

insights into relationships.” Hoffman & Howell went on to speculate that “The impression is thus gained

ENGHOFF H., New millipede genus and species from the Udzungwa Mts

of a group of organisms which have stabilized their general Gestalt whilst expressing innate genetic

variability dominantly in permutations of male reproductive structures. A young evolutionary status is

implied by the paucity of strong discontinuities in character systems and typical spectral expression of

traits.” Anybody who has tried to refer odontopygid specimens to a genus will agree that this is not easy.

Whereas the gonopods of each species are often highly characteristic, when it comes to grouping the

species, the problems are massive.

As an illustrative example, one may mention the record of Odontopyge cf. picea by Dieudonne (2014).

The record is illustrated with a very nice optical photo of the gonopods, and they do indeed look very

much like Geotypodon picea gen. et comb. nov. Examination of several specimens from among the

material recorded by Dieudonne, kindly put at my disposal by Didier VandenSpiegel (Royal Museum

for Central Africa, Tervuren, Belgium), did, however, reveal that whereas the specimens in most respects

agree with the definition of Geotypodon gen. nov. given above, they differ in a striking detail: just distal

to the posttorsal narrowing where the solenomere and the telomere begin, there is a peculiar sclerite

protruding from the main telopodite axis. The basal telomeral spine sometimes curves between this

sclerite and the main axis. Such a sclerite is absent from the true G. picea (Attems 1938: fig. 32).

Furthermore, the solenomere of these specimens has a tiny spine-like side branch near the tip, whereas

the solenomere tip is not visible on Attems’ figure and is not mentioned in the description. (As noted

above, one of the few differences between G. iringensis gen. et sp. nov. and G. multianulatus is that the

latter species has a similar small subapical spine.)

An alternative approach to odontopygid classification is clearly needed, and molecular characters are the

obvious choice. This will, however, require a huge effort. A search for “Odontopygidae” in GenBank

yields no results whatsoever, so there is scope for large-scale sampling and sequencing of well-vouchered

odontopygids.

This way, a more satisfactory classification may one day be obtained, but until then, in order to be

able to handle the amazing diversity of odontopygids, several existing genera need to be revised and

probably split into several smaller genera. This approach might seem to be at variance with the ideas

expressed by Hoffman & Howell (2012), i.e., “traditional reliance on a few obvious key-characters

(<a priori definition) instead of groupings made on the basis of overall similarity”, but at least as a

temporary solution such a splitting approach will be useful. Taking the monographic work of Kraus

(1960, 1966) as the starting point, several such splits have already been made by Demange (1981),

Frederiksen & Enghoff (2015), Hoffman (2002) and Hoffman & Howell (2012). One further split is

proposed in the present paper.

Around 400 species of Odontopygidae have been described (Enghoff 2014), but virtually every new

collection of millipedes from the Afrotropical region contains undescribed species. The number of

described odontopygid species can therefore potentially be multiplied by an unknown factor (5? 10?),

and the number of genera will - at least temporarily - also need to be multiplied, hopefully to a somewhat

smaller degree.

Acknowledgements

Thanks are due to Mogens Andersen, Peter Gravlund, Andy Jakobsen and Fine Sorensen, as well as

Frontier Tanzania, for collecting the specimens studied here, to Hans Reip for providing access to

old literature, to Nicholas Ioannou for photography, and to Didier VandenSpiegel, Royal Museum for

Central Africa, Tervuren, Belgium, for lending important comparative material.

European Journal of Taxonomy 177: 1-19 (2016)

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Manuscript received: 8 October 2015

Manuscript accepted: 26 November 2015

Published on: 26 February 2016

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.