A mountain of millipedes V: three new genera of Odontopygidae from the Udzungwa mountains, Tanzania (Diplopoda, Spirostreptida, Odontopygidae)

. Three new genera of Odontopygidae are described, all based on new species from the Udzungwa mountains, Tanzania, and all monotypic: Casuariverpa gen. nov. (type species: C. scarpa gen. et sp. nov.), Yia gen. nov. (type species: Y. geminispina gen. et sp. nov.), and Utiliverpa gen. nov. (type species: U. decapsulatrix gen. et sp. nov.). Similarities and differences between the new genera and other genera are discussed.


Introduction
This is the fifth 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  and Enghoff (2016aEnghoff ( , 2016b.
In the present contribution, three new, monotypic genera and their type species are described.

Material and methods
The material for this article comes mostly from the zoological collections of the Natural History Museum of Denmark, University of Copenhagen (ZMUC). These specimens were collected during field trips by ZMUC staff and students. A specimen from Virginia Museum of Natural History (VMNH) was also examined. All specimens are kept in 70% alcohol.
Specimens were examined in alcohol under a stereo microscope. Specimens for scanning electron microscopy (SEM) were transferred to 96% ethanol, then to acetone, air-dried, mounted on aluminium

R e s e a r c h a r t i c l e
urn:lsid:zoobank.org:pub:A9E7A041-A454-4BC1-BCF4-F1E021BFDCD5 stubs or on pieces of flexible aluminium tape and in turn mounted on stubs, coated with platinumpalladium and studied in a JEOL JSM-6335F scanning electron microscope.
As in previous articles in this series, only adult males are considered. A total of 14 adult males of the three new species were examined. Fig. 1 shows the Udzungwa localities where the new species described here were collected. Fig. 2 gives the body size (body diameter/number of podous rings) of males of the new species.
See Enghoff (2014) for the description standards used.

Other included species
None.

Diagnosis
A genus of Odontopygidae-Prionopetalini characterized by: anal valves with a well-developed dorsal spine, a similarly well-developed ventral spine and strongly protruding setiferous tubercles -male body ring seven with large ventral lobes -limbus very broad, with a simply denticulate margin -ventral postfemoral and tibial pads on some male legs -gonopod coxa simple, metaplica (mp) with a subapical basad spine (msp) -gonopod telopodite with a compact torsotope (tt) -no pretorsal or torsal spines/ processes -a pronounced posttorsal narrowing (pn) without spines -a division of the telopodite into solenomere (slm) and telomere (tlm) immediately distal to posttorsal narrowing -solenomere whiplike, with a very long accessory spine (ps) branching off near its base and with a small subdistal lobetelomere simple, with a longitudinal ridge on concave side enveloping solenomere, distally expanded, with micro-spinose fields basally and distally.

Etymology
From Latin Casuaria, "cassowary", and "verpa", penis. Refers to the shape of the solenomere tip which is somewhat reminiscent of the head profile of the Australian cassowary (Casuarius casuarius (Linnaeus, 1758)). Gender feminine.

Remarks
In the key of Kraus (1966), C. scarpa gen. et sp. nov. runs to Trichochaleponcus Attems, 1928 (couplet 21). They key character for this genus is the very broad limbus (which is, however, paralleled in certain other odontopygids, e.g., Kompsoprium firmosum Kraus, 1960(Kraus 1960 fig. 69)). Casuariverpa scarpa gen. et sp. nov. also agrees with Trichochaleponcus species in some other characters, but there are differences enough, at least vis-a-vis the type species of Trichochaleponcus, that a new genus is warranted, cf. Enghoff (2016a). Thus, Casuariverpa gen. nov. has strongly protruding setiferous tubercles and an exceptionally well-developed ventral spine on the anal valves, its gonopod coxa has no lateral process or spine at mid-length, its gonopod telopodite has no spine between torsotope and posttorsal narrowing, the solenomere has no second basal spine in addition to the very long one, and the telomere is expanded distally instead of being strongly attenuate. See Table 1.

Etymology
The name is a noun in apposition and refers to the Udzungwa Scarp Forest Reserve.
Colour. Faded; traces of a broad, light dorsal stripe.
Body ringS. Almost perfect cylinders, not vaulted; suture straight; ozopores ca three diameters behind suture.  European Journal of Taxonomy 221: 1-17 (2016) anal valveS (Fig. 3A, C). Surface, like that of preanal ring and subanal scale, coarsely rugose. Each valve with a long dorsal spine and an almost as long downwards curved ventral spine; marginal rim barely raised, setiferous tubercles strongly protruding in lateral view, but not protruding in a laterad direction.
male legS. With postfemoral and tibial ventral pads in anterior body half, except on first few leg-pairs. Body ring 7 (Fig. 3D). With large ventral lobes, much larger than usual in the family.
gonopod telopodite (Figs 3F-H, 4). Arculus ca 90°. Torsotope (tt) simple, compact, without processes, partly hidden inside coxal cavity, i.e., the part of the telopodite between arculus and torsotope extremely short. Posttorsal narrowing (pn) without processes or spines. Telopodite just distal to posttorsal narrowing dividing into solenomere and telomere. Solenomere (slm) long, whiplike, longer than telomere, with a very long accessory spine (ps) branching off near its base; tip of solenomere with a curved, triangularpointed lobe giving the tip a profile somewhat similar to that of a cassowary (see etymology). Telomere (tlm) proximally enveloping base of solenomere, distally expanded, all in all forming a roughly triangular concave plate; concavity with a longitudinal ridge (tr) delimiting a broad furrow (tf) for accommodation of solenomere; distal margin of telomere straight, gently undulate. A micro-spinose field on basal external side of telomere basis (Figs 3G, 4E), another micro-spinose field (Figs 3H, 4F) on distal part of telomere concavity.

Distribution and habitat
Known only from the two sites in and adjacent to the Udzungwa Scarp Forest Reserve, Udzungwa Mts.

Other included species
None.

Diagnosis
A genus of Odontopygidae-Prionopetalini characterized by: a metaplical shelf (ms) with a curved spine (msp) on the coxa (shared with the Chaleponcus dabagaensis group, see Enghoff 2014) -a compact torsotope (tt) -lack of pretorsal or torsal spines/processes -a pronounced posttorsal narrowing (pn) without spines -a division of the telopodite into solenomere (slm) and telomere (tlm) immediately distal to posttorsal narrowing -a pair of long spines (ts1, ts2) emerging from the base of the telomerea slender, whiplike solenomere -a very long, slender distal telomeral process (tdp) with a row of long Y-or I-shaped spikes along one edge and an extensively spinose tip.

Etymology
The name is an artificial word and refers to the row of Y-and I-shaped spikes on the gonopod telomere (Fig. 6F). To be treated as feminine.

Remarks
In the key of Kraus (1966) Y. geminispina gen. et sp. nov. runs to Odontopyge Brandt, 1841. As explained by Hoffman (1991) and Enghoff (2016a) this name has been misapplied, and species classified in Odontopyge by pre-1991 authors need to be re-allocated. This is an ongoing process, but the type species of Yia gen. nov. cannot be accommodated in any of the genera which have so far absorbed former Odontopyge species (see Enghoff 2016a). I therefore propose a new genus. None of the still orphaned ex-Odontopyge species share the diagnostic characters of Yia gen. nov., which thus remains monotypic for the time being. Geotypodon heteromodestus (Kraus, 1960) (= Haplothysanus modestus Attems, 1953, preoccupied) shares the double telopodital spine characteristic of Yia gen. nov., but differs strongly in several other characters including the structure of the telomere. A very long, very slender distal telomeral process is found in some other odontopygids, e.g., all species of Allantogonus Attems, 1912(Kraus 1960 and Prionopetalum Attems, 1909(VandenSpiegel & Pierrard 2009Enghoff 2016b), but these genera differ from Yia gen. nov. in other characters, and no other species has the row of long Y-or I-shaped spikes characteristic of this genus. The metaplical shelf and spine resemble a similar structure in the Chaleponcus dabagaensis group (Enghoff 2014), but in Yia gen. nov. the spine originates between the shelf and the apical metaplical hood, not from the shelf itself as in the C. dabagaensis group. Also the apical metaplical cucullus (cu) resembles the C. dabagensis group. Yia geminispina gen. et sp. nov. also shows specific similarities with species of the genus Trichochaleponcus Attems, 1928: the general shape of the coxa, the presence of two spines originating at the base of the telomere, and the tip of the telomere being drawn out into a slender arm. Notable differences include the limbus which is very narrow in Yia gen. nov., very broad in Trichochaleponcus, the absence of a posttorsal, pre-narrowing spine as well as of microtrichose areas on the telomere in Yia, and the unique Y-shaped spicules on the telomere tip in Yia. See also

Etymology
The name is a composite Latin noun in apposition meaning "twin spine" and refers to the pair of long spines on the telomere.  Size. Length ca 4 cm, diameter 2.6 mm, 49 podous rings, no apodous rings in front of telson.
Colour. Faded, very faint traces of a broad, light dorsal stripe.
Collum. With a marginal and a submarginal furrow.
anal valveS. Each with a long, curved dorsal spine; marginal rim slightly raised, setiferous tubercles slightly protruding in a laterad direction (i.e., on very small 'ravelins').
ozoporeS. Starting from ring 6. limBuS (Fig. 5E). With simple, slender-triangular lobes. Surface of lobes longitudinally microstriate. male legS. With postfemoral and tibial ventral pads in anterior half of body, except for first few legpairs.

Distribution and habitat
Known only from the type locality.

Other included species
None.
ENGHOFF H., Three new genera of Odontopygidae from the Udzungwa mountains A genus of Odontopygidae-Prionopetalini characterized by: a long basad metaplical spine (msp) on the anterior side of the coxa -a compact torsotope (tt) -lack of pretorsal or torsal spines/processes -a pronounced posttorsal narrowing without spines -a division of the telopodite into solenomere (slm) and telomere (tlm) immediately distal to posttorsal narrowing -a spine (ps) emerging from the base of the solenomere -a ribbonlike solenomere which is accommodated in the concavity of the telomere and which apically is divided into a long hook and a subapical pointed lobe.

Etymology
From Latin utilis: useful, and verpa: penis. Refers to the shape of the solenomere which resembles a very useful instrument: a bottle-opener. Gender feminine.

Remarks
In the key of Kraus (1966) U. decapsulatrix gen. et sp. nov. runs to Rhamphidarpoides Kraus, 1960. This genus has been a repository for a number of quite different species, but was given a more strict definition by . The new species does not fit the definition of Rhamphidarpoides s.s., nor of Raduliverpa , which was erected to accommodate several species formerly assigned to Rhamphidarpoides. I therefore propose a new genus here, but leave the question open whether some of the species "orphaned" from Rhamphidarpoides by  might belong here.

Etymology
The name is a Latin noun meaning "remover of capsules", cf. etymology of genus.
Colour. Specimen faded, traces of a broad, light dorsal stripe. Colour according to the collector's field notes: "dark, shiny black with brown stripe length of back; legs pale".
Collum. With a marginal and a submarginal furrow.
Body ringS. Almost perfect cylinders, not vaulted; suture straight; ozopores ca three diameters behind suture.
anal valveS. Each with a short, curved dorsal spine and a small ventral denticle; marginal rim slightly raised, setiferous tubercles hardly protruding.

Distribution and habitat
Known only from the type locality. Habitat: forest (under log).

Coexisting species
No other odontopygid species were found in the same sample as the unique holotype, but three are known from Mwanihana Forest Reserve: Chaleponcus mwanihanensis Enghoff, 2014, Aquattuor major A. submajor Enghoff, 2015.

Discussion
In a previous paper (Enghoff 2016a), I argued that due to the bewildering confusion about genus concepts in Odontopygidae and to the absence of any well-supported hypotheses of relationships within the family, a 'splitter' approach will probably serve best, at least at the present stage. This means that a considerable number of narrowly circumscribed genera need to be defined, an agenda already adhered to by, e.g.,  and Enghoff (2016a). In the same spirit three new monotypic genera are described in the present paper. Including these, the number of described odontopygid species from the Udzungwa mountains is 33. A few additional species, belonging to (relatively) well-defined, known genera, are known to exist, and ongoing and planned field-work in the Udzungwas will certainly reveal many more.