European Journal of Taxonomy 83: 1-36

http://dx.doi.org/10.5852/eit.2014.83

www. europeani ournaloftaxonomv. eu

2014 • Mesibov R.

ISSN 2118-9773

This work is licensed under a Creative Commons Attribution 3,0 License.

Research article

urn:lsid:zoobank.org:pub:049F326B-9460-4038-BB21-9DA36F79812F

The millipede genus Solaenodolichopus Verhoeff, 1924 (Diplopoda,

Polydesm da, Paradoxosomatidae). 1. New genus diagnosis and

redescriptions of named species

Robert MESIBOV

Queen Victoria Museum and Art Gallery, Faunceston, Tasmania, Australia 7250

Email: [email protected]

urn:lsid:zoobank.org:author:24BA85AE-1266-494F-9DE5-EEF3C9815269

Abstract. Solaenodolichopus Verhoeff, 1924 is redefined to include S. pruvoti (Brolemann, 1931), S.

rubriventris Verhoeffi 1928, S. sulcatus (Verhoeff, 1928), S. teres (Verhoeff, 1924), S. vittatus (Verhoeff,

1924) and S. walesius Verhoeff, 1928, each of which is redescribed. Lectotypes are designated for

S. sulcatus , S. teres, S. vittatus and S. walesius. Parw ale soma Verhoeff, 1937 is synonymised with

Solaenodolichopus and S. vittatus dorsalis (Verhoeff, 1924) with S. vittatus vittatus (Verhoeff, 1924).

Keywords. Millipede, Paradoxosomatidae, Queensland, New South Wales, Australia.

Mesibov R. 2014. The millipede genus Solaenodolichopus Verhoeff, 1924 (Diplopoda, Polydesmida, Paradoxo¬

somatidae). 1. New genus diagnosis and redescriptions of named species. European Journal of Taxonomy 83: 1-36.

http://dx.doi.org/10.5852/eit.2014.83

Introduction

Solaenodolichopus Verhoeff, 1924 is an eastern Australian genus of Paradoxosomatidae with a

distinctive gonopod structure and no obvious paranota on diplosegments. There are six named and many

undescribed species native to Queensland (Mesibov 2008) and New South Wales (Car 2009). One of the

named species, S. pruvoti (Brolemann, 1931), was first described from New Caledonia and has recently

been reported from Western Australia (Car et al. 2013); more records are reported here from southeast

Queensland, where S. pruvoti is likely to be native. Records of S. pruvoti from South Australia (Jeekel

1982, 2002) may be based on a misidentification (see below).

In this paper I review the history of the Solaenodolichopus genus concept, propose a new diagnosis,

redescribe the six named species, designate several lectotypes, accept two previously proposed

synonymies and propose two new synonymies. New species of Solaenodolichopus will be treated in

future papers in this series.

Materials and Methods

‘Male’ and ‘female’ in the text refer to adult individuals. All specimens are stored in 70-80% ethanol in

their respective repositories.

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European Journal of Taxonomy 83 : 1-36 ( 2014 )

The photomicrographs in Fig. 11C and 11D were taken using an eyepiece video camera mounted on an

Amscope binocular microscope. Other photomicrographs were taken with a Canon EOS 1000D digital

SLR camera mounted on a Nikon SMZ800 binocular dissecting microscope equipped with a beam

splitter. Measurements were made to the nearest 0.1 mm with the same microscope using an eyepiece

grid and a reference scale. Most photomicrographs used in the figures are focus-stacked composites

prepared with Zerene Stacker 1.04 software. Preliminary gonopod drawings were traced from printed

copies of focus-stacked photomicrographs; details were added and corrected by reference to the actual

gonopod. SEM images were acquired digitally with a FEI Quanta 600 ESEM at low vacuum using

uncoated specimens which were air-dried, examined and returned to alcohol. Maps were generated

using Arc View GIS 3.2. All figures were prepared using GIMP 2.8 image editing software; backgrounds

in some photomicrographs have been edited to remove distracting highlights and artifacts.

Latitude/longitude figures are given with the WGS84 datum together with my estimate of the spatial

uncertainty (Darwin Core coordinateUncertaintylnMeters). Data in square brackets are my additions;

latitude/longitude figures not in square brackets are based on label data or the respective museum

collection databases.

Details of specimen lots of the species redescribed in this paper are tabulated as an Excell file in

Supplement E

Images of Solaenodolichopus types as alcohol specimens and slide mounts, including their labels, are

compiled in a PDF file in Supplement 2.

Abbreviations

AM

NHRS

NSW

QLD

QM

SAM

ZMB

ZSM

Australian Museum, Sydney, Australia

Naturhistoriska Riksmuseet, Stockholm, Sweden

New South Wales, Australia

Queensland, Australia

Queensland Museum, Brisbane, Australia

South Australian Museum, Adelaide, Australia

Museum far Naturkunde, Berlin, Germany

Zoologische Staatssammlung Miinchen, Munich, Germany

Results

Order Polydesmida Pocock, 1887

Suborder Strongylosomatidea Brolemann, 1916

Family Paradoxosomatidae Daday, 1889

Subfamily Australiosomatinae Brolemann, 1916

Tribe Antichiropodini Brolemann, 1916

Solaenodolichopus Verhoeff, 1924

Solanodolichopus Verhoeff, 1924: 10 (in key as subgenus of Antichiropus), 11 (in key as subgenus of

Antichiropus ), 19 (first description).

Paraulacoporus Verhoeff* 1928: 88, 90 (in key), 91 (in key), 93. [Type species Paraulacoporus sulcatus

Verhoeff, 1928, by monotypy.]

Parwalesoma Verhoeff, 1937: 139 (first description). [New synonymy. Type species Parwalesoma

castaneum Verhoeff, 1937, by monotypy.]

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MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Solaenodolichopus - Attems 1926: 143 (in key as subgenus of Antichiropus)', 1937: 260 (synonymised

with Aulacoporus). — Verhoeff 1932: 1574 (as subgenus of Antichiropus ), 1595 (in key), 1597 (in

key). —Jeelcel 1965: 8; 1968: 29 (as synonym of Aulacoporus)', 1971: 233 (type species designated);

2000: 37 (in key), 4CM2; 2003: 42. —Hoffman 1980: 166 (as synonym of Aulacoporus). —Nguyen

& Sierwald 2013: 1159.

Solanodolichopus - Verhoeff 1928: 88 (raised to genus), 90 (in key), 91 (in key), 94. — Brolemann

1931: 299. —Verhoeff 1937: 139. —Verhoeff 1941: 8 (in key).

Par aulacoporus - Attems 1929: 258. — Brolemann 1931: 295, 298. — Verhoeff 1932: 1595 (in key),

1598 (in key). — Attems 1937: 260 (synonymised with Aulacoporus). — Verhoeff 1941: 8 (in key).

— Jeelcel 1965: 8. — Jeelcel 1968: 29 (as synonym of Aulacoporus). — Jeelcel 1971: 230 (in genus

catalogue). — Hoffman 1980: 166 (as synonym of Aulacoporus). — Jeekel 2000: 40, 42 (synonymised

with Solaenodolichopus). —Nguyen & Sierwald 2013: 1159 (as synonym of Solaenodolichopus).

Parwalesoma - Attems 1940: 549 (Verhoeff s description reworded). — Verhoeff 1941: 8 (in key). —

Jeekel 1965: 8. — Jeekel 1968: 29, 126. — Jeekel 1971: 231. —Hoffman 1980: 166. — Jeekel2000: 37

(in key), 40-42. — Jeekel 2003: 42. — Nguyen & Sierwald 2013: 1157.

Type species

Antichiropus (Solaenodolichopus) teres Verhoeff, 1924, by subsequent designation.

Other assigned species

S. pruvoti (Brolemann, 1931), S. rubriventris Verhoeff, 1928, S. sulcatus (Verhoeff, 1928), S. vittatus

(Verhoeff, 1924) and A walesius Verhoeff, 1928.

History of Solaenodolichopus

The sometimes confusing development of the Solaenodolichopus concept over the past 90 years has

involved five genus names and four diplopodologists.

Besides Solaenodolichopus , some of the species redescribed in this paper have also been assigned

to Antichiropus Attems, 1911, Aulacoporus Verhoeff, 1924, Par aula cop or us Verhoeff, 1928 and

Parwalesoma Verhoeffi 1937.

The four diplopodologists were the Austrian Carl Attems (1868-1952, in full, Carl Aug. Graf [Count]

Attems-Petzenstein), the German Karl W ilh e lm Verhoeff (1867-1944), the Frenc hm an Henry Wilfred

Brolemann (1860-1933) and the Dutchman Casimir Albrecht Willem Jeekel (1922-2010). I use the

spelling ‘Brolemann’ below rather than ‘Brolemann’, following that author’s practice in the 1931

publication cited in this paper. Brolemann dropped the umlaut in his name from 1920 onwards (J.-J.

Geoffroy, in litt., 30 Apr. 2009).

All four specialists viewed the polydesmidan gonopod as a modified walking leg on which it was

possible to recognise podomere homologs. Their differing opinions on homology are mentioned only

briefly below in the context of a chronology of Solaenodolichopus. My purpose here is to try to explain

the reasoning behind particular generic assignments in the past, not to thoroughly review an enduring

and still-unresolved controversy.

Solaenodolichopus was first proposed by Verhoeff (1924) as a subgenus of Antichiropus Attems, 1911.

Attems (1911) had described seven new species from Western Australia with gonopod telopodites divided

into three main parts: a small, densely setose, basal portion ( Femur ); a long, cylindrical, slender, central

portion {Tibia) tipped with short prongs ( Zacken ); and a long, thin, coiled, apical portion {Kanalast)

carrying the prostatic groove and its terminal opening. Attems assumed that the tarsus of the modified

leg had fused with the tibia, and that the distal portion of the tibia was its tarsal section ( Tarsalabschnitt ).

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European Journal of Taxonomy 83: 1-36 (2014)

In his view, the first joint of this tarsal section consisted of the Kanalast and a prong-like process ‘a’ at the

end of the tibia and close to the prostatic groove. Attems suggested that another tibial process, ‘c’, was

a remnant of a second tarsal joint. (For a discussion of the Antichiropus gonopod and its components,

see Car et al. 2013.)

Verhoeff (1924) described the telopodites of the new central Queensland species teres and the southern

Queensland vittatus (the latter divided into two subspecies) as likewise having a small, densely

setose, basal portion (. Praefemur ); a long, cylindrical, slender, central portion (Femur) tipped with

short accessory branches (. Nebendste ); and a long, terminal process carrying the prostatic groove and

its opening. Verhoeff regarded the latter as an evolutionarily novel structure (Neubildung) he called a

Solanomerit, which he had recognised in many other Polydesmida and which had no homologue on

a walking leg; in what follows I will use the modern term ‘solenomere’. The two Queensland species

were placed in Antichiropus (Solaenodolichopus) not because this arrangement was definitive (wrote

Verhoeff), but because the two species seemed more closely related to those in Antichiropus than to any

other species he had previously examined. The gonopod telopodites were fundamentally similar and the

first male legs (bearing a short, rounded, ventral, femoral process) were almost identical.

According to Verhoeff (1924), the polydesmidan solenomere was primitively protected by a long

tibiotarsus, but there were four secondary developments away from this pattern. In one development

(Verhoeff s case 1), the tibiotarsus was reduced so much that the solenomere became the free end of the

gonopod, as in Antichiropus. The tibiotarsus had either disappeared completely in Antichiropus , or had

been reduced to one of the small accessory branches at the end of the femur (Attems’ Zacken). In another

development (case 4), the tibiotarsus and solenomere fused into a single process. The two developments

could overlap (Verhoeff 1924: 6, “Die Falle N. 1 und 4 gehen fibrigens in einander fiber”). In the new

Queensland species, Verhoeff believed he could detect a fusion in which the solenomere had dominated,

so that the tibiotarsus was reduced to a small subterminal tooth on the teres solenomere and a larger but

likewise distal process on the solenomere in vittatus.

In the same 1924 paper, Verhoeff proposed Aulacoporus as a new genus for three northern Queensland

species. Aulacoporus was distinguished from Antichiropus (Solaenodolichopus) in two ways. First, the

prostatic groove opened at the extreme tip of the solenomere in Aulacoporus , whereas in Antichiropus

(Solaenodolichopus) the opening was carried on a near-apical process of the solenomere. The

solenomere and tibiotarsus had fused in Aulacoporus, but the tibiotarsus had completely disappeared in

one Aulacoporus species and in the other two was reduced to a distal bulge on the solenomere.

To explain Verhoeff’s second diagnostic difference between Aulacoporus and Antichiropus

(iSolaenodolichopus ), I need to digress briefly.

Polydesmidan species have been classified, in part, on the structure of the paranota, which are lateral

extensions of the body wall. In many families the paranota are wide, dorsoventrally flattened and more or

less horizontal. In some families there are species with no detectable paranota, at least on diplosegments,

i.e. the body rings are simple circles in cross-section. Between these two extremes, paranota are often

said to be ‘reduced’. Verhoeff (1924) attempted to refine the description of paranota for classification

purposes. Instead of Kiel (keel, carina) for paranota in general, Verhoeff proposed Seitenwiilste (lateral

bulges) for all lateral swellings and extensions. In species with wide, dorsoventrally flattened, wing-like

Seitenwiilste , the term Seitenfliigel (lateral wings) would be appropriate. Each Seitenwulst was often

accompanied medially by a narrow, dorsal, longitudinal furrow which Verhoeff called a Seitenfurche

(lateral groove). Species with no Seitenwiilste at all might or might not have Seitenfurchen.

4

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Attems (1937) used these terms slightly differently. He observed that well-developed paranota, i.e.,

Seitenfliigel , often had swollen lateral margins. In some species the paranota had narrowed so much that

only these marginal thickenings were still present, in which case one could speak of Seitenwiilste , and in

species without Seitenwiilste there might still remain the medial, demarcating grooves, the Seitenfurchen.

Both Attems and Verhoeff understood these variations as an evolutionary series in which primitively

wide paranota were progressively reduced to bulges and, before disappearing entirely, to short, fine,

lateral grooves on the metazonites.

The Aulacoporus and Antichiropus (*Solaenodolichopus ) species described by Verhoeff had reduced

but detectable paranota on the haplosegments, i.e. rings 2-4. Neither group had Seitenwiilste on the

diplosegments, but there were distinct Seitenfurchen in Aulacoporus and none at all in Antichiropus

( Solaenodolichopus ).

Attems (1926) accepted both Aulacoporus and Solaenodolichopus (as a subgenus of Antichiropus ),

but reiterated his view that beyond the coxa on the gonopod were only a femur, tibia and tarsus.

Verhoeff s Solanomerit was simply a tibial extension ( Tibialfortsatz ). In a key to paradoxosomatid

genera (‘Strongylosomidae’), Attems (1926) separated Aulacoporus (and other genera) having a simple

telopodite without side branches or a tarsal section, from Antichiropus (and other genera) having a multi-

tipped telopodite. The key listed Antichiropus as having 1-3 short processes, probably tarsal remnants,

near the base of the tibial extension (i.e., the solenomere).

Verhoeff (1928) raised Solaenodolichopus to a genus and added two new species from NSW,

rubriventris and walesius. He also proposed Par aulacoporus for another new NSW species, sulcatus. In

Par aulacoporus as in Aulacoporus there were short Seitenfurchen on most diplosegments, although the

grooves were indistinct on rings 15-19. Unlik q Aulacoporus but like Solaenodolichopus , Paraulacoporus

was characterised by a prostatic groove opening proximal to the solenomere tip. Unlike the situation in

Solaenodolichopus , the two accessory branches at the end of the femur in Paraulacoporus were of equal

size, and the process lateral to the prostatic groove opening on the solenomere was forked. Verhoeff

described this forked process as a ‘tarsal branch’.

Brolemann (1931) assigned a new New Caledonian species, pruvoti, to Paraulacoporus , although he

observed no lateral groove (sillon) past ring 5. Combining the ideas of Attems and Verhoeff, Brolemann

described the pruvoti telopodite as consisting of a short, densely setose, basal femoroid (femoroide ), a

long, slender tibia and a terminal seminal branch (rameau seminale). He regarded the short, posteromedial

process at the end of the tibia as a reduced tarsus separate from the seminal branch, and rejected Verhoeffs

opinion that the seminal branch in P. sulcatus was a fusion of solenomere and tibiotarsus. Brolemann

also objected to the placement of Solaenodolichopus within Antichiropus by Verhoeff and Attems. He

suggested instead that Paraulacoporus and Solaenodolichopus could be grouped together because both

had an erect, expanded, more or less divided seminal branch, not a long, coiled strap as in Antichiropus.

Verhoeff (1932) offered simple new keys to the known Australian paradoxosomatids (‘Strongyloso¬

midae’) based either on trunk details or gonopod structure. In the latter key, Aulacoporus , Paraulaco¬

porus, Solaenodolichopus and Walesoma Verhoeff, 1928 were grouped as genera in which a long

solenomere led from the end of the femur, which also might have 1-2 accessory branches at its apex.

Aulacoporus and Walesoma were paired as genera in which the opening of the prostatic groove was at

the extreme end of the telopodite, with no accessory process either proximal or distal to the opening.

Paraulacoporus and Solaenodolichopus were paired as genera in which the opening of the prostatic

groove was proximal to the extreme end of the telopodite, with accessory processes both proximal and

distal to the opening.

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European Journal of Taxonomy 83: 1-36 (2014)

Attems (1937) revised his nomenclature of sections of the gonopod telopodite, renaming femur, tibia

and tarsus as prefemur, femur and tibiotarsus. The process carrying the prostatic groove ( Samenrinne )

and its opening was the Rinnenast (groove-branch) and was a process of the femur. More importantly,

in 1937 Attems synonymised both Paraulacoporus and Solaenodolichopus under Aulacoporus. He

described the gonopod and paranota in the expanded genus as follows (Attems 1937: 260-261, my

translation): “Gonopod femur long to very long and slender, at the end with one or no side-branch.

Rinnenast very large and broad, with one, sometimes two side-teeth. The tibiotarsus is reduced to a

small spine. Seitenwiilste wealdy developed, low, round, sometimes only present on anterior segments.

Tongitudinal grooves either present on most segments or lacking from segments 5 or 6 onwards”. In

his key to Aulacoporus species, Attems separated the former Aulacoporus species from the former

Paraulacoporus and Solaenodolichopus species simply on the presence or absence of Seitenfurchen on

most rings.

Verhoeff (1937) described the new species castaneum from NSW and assigned it to a new genus,

Parwalesoma. He noted that Parwalesoma was close to both Aulacoporus and Solaenodolichopus ,

but differed from the former genus in lacking Seitenfurchen on most rings. Parwalesoma and

Solaenodolichopus differed in a somewhat sharper demarcation between femur and solenomere, and in

the solenomere tip in P. castaneum being spoon-shaped. Parwalesoma castaneum Verhoeff, 1937 was

in fact Solaenodolichopus walesius Verhoeff, 1928 redescribed from new material (see below under S.

walesius ), but it was another 60+ years before Verhoeffs confusion was detected by Jeekel (2000; see

below).

In his last contribution to the Solaenodolichopus story, Verhoeff (1941) unwittingly redescribed

Brolemarm’s S. pruvoti (from New Caledonia) as Solaenodolichopus annulatus (from Brisbane). He

also offered a key to genera in which he separated the taxa lumped by Attems (1937). The relevant

portion of the key is as follows (Verhoeff 1941: 8, my translation with Verhoeffs terminology updated):

c) Solenomere divided at the end into two branches, the medial one extending as three protuberances,

of which the middle contains the opening of the prostatic groove, the proximal one a bend in the

prostatic groove. Trunk without Seitenwiilste but with Seitenfurchen . Paraulacoporus Verh.

d) Prostatic groove without bend before its end .e, f

e) Solenomere before its end with a spoon-shaped concavity. Trunk with neither Seitenwiilste nor

Seitenfurchen . Parwalesoma Verh.

f) Solenomere without this concavity .g, h

g) End of the solenomere drawn out into two closely standing tips, of which the more proximal

contains the opening of the prostatic groove. Trunk with neither Seitenwiilste nor Seitenfurchen

. Solaenodolichopus Verh.

h) Opening of the prostatic groove in a simple, apical tip.i, j

i) Solenomere significantly shorter than femur, its base enlarged, its end hooked. Trunk without

Seitenwiilste but with Seitenfurchen . Aulacoporus Verh.

j) Solenomere and femur about equally long, the solenomere base not swollen. Trunk with weak,

posteriorly rounded Seitenwiilste and Seitenfurchen . Walesoma Verh.

Reviewing the Australian Antichiropodini, Jeekel (1968) accepted Attems’ (1937) synonymies and

Verhoeffs Parwalesoma. Thirty-two years later, he confessed that he regarded Parwalesoma as another

Aulacoporus synonym, but had not published this synonymy because the Parwalesoma type species,

P. castaneum Verhoeff, 1937, would have become a junior homonym of the Aulacoporus type species,

A. castaneus Verhoeff, 1924. “As the broad concept of Aulacoporus was intended to be a temporary

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MESIBOV R., Redescriptions of Solaenodolichopus millipedes

solution anyway, the introduction of a new name for the type-species of Parwalesoma seemed just too

radical” (Jeekel 2000: 40).

Jeekel (2000) then reinstated Solaenodolichopus , leaving within it S. teres , S. vittatus and S. vittatus

dorsalis and transferring to it the Paraulacoporus species P. sulcatus and P. pruvoti. He also recognised

S. annulatus as a synonym of pruvoti. Solaenodolichopus now included species with and without

Seitenfurchen, and with considerable variation in the size and position of the processes at the end of

the telopodite femur. “The species of this genus are characterised by the solenomerite of the gonopods

having a pre-apical latero-distal process paralleling the apex proper. This process distinguishes the genus

from the others” (Jeekel 2000: 41).

One of those other genera was Aulacoporus and its three original species, which Jeekel (2000) regarded

as distinct not because the solenomere was simple with the prostatic groove opening at its tip, but

because the gonopod apex curved strongly cephalad and bore a large lobe laterally - a lobe which Jeekel

presumed was the tibiotarsus.

The other genus Jeekel (2000) distinguished from Solaenodolichopus was Parwalesoma. He recognised

the type species P. castaneum as a junior subjective synonym of S. walesius , thus establishing P. walesium

(Verhoeff, 1928). He also transferred S. rubriventris to Parwalesoma , noting that “In this genus the

gonopods are lacking a pre-apical process on the solenomerite, but otherwise are quite similar to those

of Solaenodolichopus ” (Jeekel 2000: 42).

Jeekel’s distinction between Parwalesoma and Solaenodolichopus is questionable in the case of S.

walesius (see the taxonomic notes following the S. walesius redescription, below), and for that reason

I am making Parwalesoma a junior subjective synonym of Solaenodolichopus. This action broadens

the circumscription of Solaenodolichopus and might allow it to be subsumed under Aulacoporus , as

favoured by Attems (1937). However, after examining late 20th-century specimens of Aulacoporus

castaneus Verhoeff, 1924, A. yarrabahnus Verhoeff, 1924 and three undescribed Aulacoporus spp., all

from a small area in the Queensland Wet Tropics, I agree with Jeekel (2000) that this genus is best kept

separate from Solaenodolichopus. All Aulacoporus forms have prominent lateral grooves and slight

paranotal swellings on all diplosegments, and in four of the six Aulacoporus forms the solenomere

curves anteriorly. None of the Solaenodolichopus I have examined (as diagnosed above) has either

prominent lateral grooves on all diplosegments or an anteriorly curving solenomere.

Diagnosis

Australian Paradoxosomatidae without distinct paranota on posterior rings, and with a gonopod

telopodite with a small, densely setose prefemur and a more or less straight, slender and cylindrical

femorite ending in an unbranched medial process, an unbranched lateral process and a much larger

solenomere with a flattened tip on which the prostatic groove usually opens medially.

Description

Male with vertex bare, frons and clypeus sparsely setose; vertigial sulcus distinct, ending anteriorly just

above level of antennal sockets; post-antennal groove shallow; antennal sockets separated by 1.0-1. IX

a socket diameter. Ante nn a filiform (Fig. 1A), reaching dorsally to rear of ring 2; ante nn omere with

relative lengths typically (2,3,4) > (5,6), the differences small, and maximum distal widths subequal

or with 6 very slightly wider. Head and collum about equal in width in dorsal view, about as wide as

rings 2-4 or a little narrower; diplosegments a little wider, slightly increasing in width to midbody,

width decreasing progressively on last rings. Collum D-shaped in dorsal view; anterior margin straight

medially, gently curving laterally; posterior margin straight or very slightly emarginate medially; collum

corner broadly rounded (Fig. 1A). Ring 2 paranotum (Fig. 1A) a slightly thickened, overhanging ridge.

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European Journal of Taxonomy 83: 1-36 (2014)

nearly straight and more or less longitudinal, corners rounded, set at ca. 1/4 ring height or a little less,

below level of collum corner. Rings 3 and 4 paranota (Fig. 1A) at about level of collum corner, reduced

to shallow furrows curving upwards anteriorly. Pleural keels present on rings 2-4 as slight thickenings

with upwardly curving margins. Diplosegments without paranotal thickenings, rings circular in cross-

section. Prozonites and metazonites smooth, bare; waist short, shallow, with suture at anterior edge,

sculptured indistinctly as variably low, rounded, longitudinal ridges; limbus a short, thin, continuous

sheet. Pore formula normal; ozopores small, round, flush with lateral surface of ring, set at mid-height

Fig. 1. — A-B. Solaenodolichopus walesius Verhoeff, 1928. A. Lectotype AM KS.76508, right

anterior view. B. $ ex AM KS. 16152, midbody spiracles, ventral view, anterior to left. — C, E. S.

vittatus (Verhoeff, 1924). C. Paralectotype §, NHRS KAS1000000007, midbody spiracles, ventral

view, anterior to left (image right-left reversed for ease of comparison with IB). E. $ ex QM S74828,

anterior leg tip with scopula, tarsus at left. — D. S. teres (Verhoeff, 1924), lectotype $ slide mount,

NHRS KAS 1000000005, legpair 1. — F. S. pruvoti (Brolemann, 1931), $ ex QM S74856, epiproct,

posterior and slightly ventral view. Scale bars: A = 2.5 mm, B-C = 0.5 mm, D = 1 mm, E = 0.25 mm, F

= 0.1 mm

8

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

and mid-length on metazonite, in line with transverse furrow (when present). Diplosegment spiracles

(Fig. 1B-C) just above and anterior to leg bases, more or less egg-shaped with smaller end ventral,

anterior spiracle larger and extended anteroventrally; spiracles with thin, short rim; spiracular filter

slightly emergent, forming rounded fold in inverted, tight U-shape in spiracular opening (Fig. 1B-C).

Midbody stemites about as long as wide, sparsely setose; cross impressions wide, shallow, transverse

impressions a little narrower and deeper than longitudinal. Feg 1 (Fig. ID) with small, bluntly rounded,

ventral, femoral process curving slightly distally, reaching to ca. 1/2 length of femur on ventral side.

Midbody legs with relative podomere lengths femur > prefemur > (postfemur, tibia, tarsus) or tarsus very

slightly longer than postfemur and tibia; femur 1.7-1.8X as long as tarsus; prefemora a little swollen

dorsally. Dense brush (scopula) (Fig. IE) of straight, finely pointed setae ventrally on tarsus and distal

portion of tibia of most legs. Epiproct extending a little past anal valves, tapering in dorsal view with

sides more or less straight, ca. 1/4-1/3 maximum width of preanal ring at apex; apex truncate, corners

produced posteriorly as small, rounded bumps (Fig. IF); spinnerets in rectangular array, wider than

long. Hypoproct subtrapezoidal to broadly paraboloid. Gonopore small, round, opening ventrally on

slight distomedial swelling of leg 2 coxa. Sternal lamella (Fig. 2) wide, ca. 80-90% of width between

leg 4 bases, perpendicular to ring or slightly tilted anteriorly, with sparse long setae on posterior surface

and dense, short setae on anterodistal surface. Aperture ca. 1/3 of ring 7 prozonite width, rounded-

rectangular and tilted slightly posteriorly; prominent, rounded, transverse swelling just anterolateral to

aperture on either side (Fig. 3A).

Female very similar to male but with legs proportionally shorter, prefemora not swollen; epigyne ca.

1/6-1/5 ring width, ends slightly raised and cradling leg 2 coxae, centre slightly raised in small, rounded

triangle (Fig. 3C).

A

b 4 7"

c

D

V

E

F

- /***< ,

A *

Fig. 2. Posterior views of sternal lamella in <$<$. — A. Solaenodolichopus pruvoti (Brolemann, 1931),

QM S74856. — B. S. rubriventris Verhoeff, 1928, QM S74690. — C. S. sulcatus (Verhoeff, 1928),

lectotype, AM KS.76503. — D. S. teres (Verhoeff, 1924), lectotype in alcohol, ZSM A20052188. — E.

S. vittatus (Verhoeff, 1924), QM S74829. — F. S. walesius Verhoeff, 1928, lectotype, AM KS.76508.

Scale bars: 0.5 mm.

9

European Journal of Taxonomy 83: 1-36 (2014)

Species-diagnostic characters

Adults of the six named Solaenodolichopus species conform to the genus description above. Omitted

from the genus description are diagnostic characters reported in the descriptions of individual species,

below. Besides gonopod structure, these species-diagnostic characters are: colour, maximum midbody

width, transverse furrow distinctness, longitudinal furrow details for diplosegments, distribution of

scopulae on legs, sternal lamella details, leg spacing on rings 6 and 8, shape of the anterior margin of the

gonopod aperture, and leg 2 coxa details in females.

Fig. 3. — A-B. Solaenodolichopus rubriventris Verhoeff, 1928, QM S74690. A. Right ventrolateral

view of ring 7 showing swelling (arrow) just anterolateral to aperture. B. Ventral view of aperture

showing gently convex extensions (arrows) on either side of medial extension on anterior margin of

aperture. — C. S. walesius Verhoeff, 1928, 5, AM KS. 16152, posteroventral view of epigyne showing

raised end (arrow). — D. S. teres (Verhoeff, 1924), paralectotype §, NHRS KAS1000000007, left

ventrolateral view of anterior rings showing distolateral process on leg 2 coxa (arrow) just medial to

raised end of epigyne. Scale bars: 1 mm.

10

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

In the species descriptions I use the terms ‘pre femur’, ‘femorite’ and ‘solenomere’ in agreement with

the usage of Car et al. (2013: 84) for Antichiropus; the terms are not intended to be implied statements

of homology.

A brief dichotomous key to the six named Solaenodolichopus species follows. The key will need major

revision as the many undescribed species are named and documented in future papers in this series.

1. Body pale yellow or light brown with narrow dark brown rings at the rear of each metazonite

. S. pruvoti (Brolemann, 1931)

- Body more or less uniformly dark in colour, with or without a lighter, medial dorsal, longitudinal

band .2

2. Longitudinal furrows laterally at level of ozopore on diplosegments to about ring 15, medial and

lateral femorite processes obvious, spine-like, about equal in length. S. sulcatus (Verhoeff, 1928)

- No longitudinal furrows laterally on any diplosegment, medial and lateral femorite processes

greatly different in length or both inconspicuously small .3

3. Lateral femorite process much larger than medial . S. walesius Verhoeff, 1928

- Medial femorite process much larger than lateral, or both femorite processes inconspicuously small

.4

4. Distal 1/3 of femorite bent at ca. 45°, femorite processes inconspicuously small.

. S. teres (Verhoeff, 1924)

- Lemorite straight, medial femorite process large and obvious.5

5. Solenomere with several spine-like processes in basal half, no large indentation in distal half

. S. rubriventris Verhoeff, 1928

- Solenomere without spine-like process in basal half, with large, U-shaped indentation in distal

half . S. vittatus (Verhoeff, 1924)

Solaenodolichopus pruvoti (Brolemann, 1931)

Tigs 1L, 2A, 4A-B, 5, 6A-B, 7, 10A, 11D

Paraulacoporus Pruvoti Brolemann, 1931: 295 (first description); figs 29-32 (p. 297), 33-35 (p. 299).

Solanodolichopus annulatus Verhoeff, 1941: 11 (first description); figs 1-4 (p. 23).

Solaenodolichopus pruvoti - Jeekel 2000: 40 (new combination), 42, 46. — Nguyen & Sierwald 2013:

1159.

Solaenodolichopus annulatus - Jeekel 2002: 60 (as synonym of Solaenodolichopus pruvoti ).

Solanodolichopus annulatus - Nguyen & Sierwald 2013: 1159 (as synonym of Solaenodolichopus

pruvoti).

Aulacoporus pruvoti - Attems 1937: 261 (new combination), 264 (Brolemann’s 1931 description

reworded); fig. 331 (p. 265; same as fig. 32 in Brolemann 1931). —Jeekel 1968: 18, 29; 1981: 47; 1985:

34.

Aulacoporus annulatus - Jeekel 1968: 20,29; 1981: 49; 2000: 41 (synonymised with Solaenodolichopus

pruvoti ), 42.

non? Solaenodolichopus pruvoti - Jeekel 2002: 60 (partial redescription from new material), 77; fig. 1

(P- 62).

non? Aulacoporus pruvoti - Jeekel 1982: 124 (redescription from new material); figs 2, 3 (p. 126).

11

European Journal of Taxonomy 83: 1-36 (2014)

Material examined

Syntypes of S. pruvoti

At least 1 male and 1 female, Dumbea, New Caledonia [22°09’ S, 166°27’ E, ±5 km], date unknown, A.

Pravot-Fol; specimens not located.

Syntypes of S. annulatus

At least 1 male, Brisbane, QLD [27°28’ S, 153°0r E, ±10 km], Dec. 1936, J. Mauritzon; specimens not

located.

Other material

QUEENSLAND: 1 Mt Glorious [27°20’ S, 152°46’ E, ±2 Ion], Oct. 1970, H. Williams, QM S5944;

1 f, Mt Nebo [27°23’ S, 152°47’ E, ±2 km], 19 Nov. 1978, A. Rozefelds, pitfall, wet sclerophyll, QM

Fig. 4. — A-B. Solaenodolichopuspruvoti (Brolemann, 1931), QM S74856. Dorsal (A) and lateral (B)

views of midbody rings of <$. — C-I. S. vittatus (Verhoeff, 1924). Dorsal (C-F) and lateral (G-I) views

of midbody rings off; QM S74767 (C, H), QM S74855 (D), QM S74836 (E, G), QM S74876 (F), QM

S5946 (I). Scale bars: 2 mm.

12

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

S5945; 2 SS, 1 ?, Camira, 27°37’ S, 152°55’ E [±2 km], 30 Jul.-Oct. 1990, R. Raven, open forest,

pitfall, QM S19631; 1 Mt Glorious barracks, 27° 19’54” S, 152°45’12” E [±500 m], 660 m a.s.l., 7 Dec.

1991-6 Mar. 1992, G. Monteith, rainforest, pitfall±intercept, QM S74833; 1 Mt Mee, 27°02’53” S,

152°40’49” E [±500 m], 300 m a.s.l., 3 Mar.-12 Apr. 1992, D. Cook, rainforest, intercept, QM S74834;

2 Stony Creek via Samford, 27°20 , 20” S, 152°47 , 52” E [±500 m], 200 m a.s.l., 22 Oct. 1994-2

Feb. 1995, H. Janetzki and G. Monteith, rainforest, pitfall, QM S74835; 1 1 $, Camerons Scrub,

knoll top, 27°30’27” S, 152°43 , 40” E [±500 m], 90 m a.s.l., 15 Sep.-ll Nov. 1998, G. Monteith, D.

Cook and G. Thompson, vine scrub, pitfall, sample 7380, QM S74841; 3 SS, 1 ?, same details but 11

Nov. 1998-13 Jan. 1999, sample 7557, QM S74856; 1 same details but Camerons Scrub, top ridge.

Fig. 5. Solaenodolichopuspruvoti (Brolemann, 1931). A. Anterior lateral view showing reach of gonopod

telopodites, QM S74856. B. Posterior view of gonopods, WAM T52345. C-D. Lateral (C) and posterior

(D) views of leg 2 coxal process in QM S74857. Scale bars: A-B = 2 mm, C-D = 1 mm.

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European Journal of Taxonomy 83: 1-36 (2014)

27 o 30’29” S, 152°43’54” E [±500 m], 160 m a.s.l., 21 Sep.-ll Nov. 1998, sample 7381, QM S74842;

I S, 1 5, same details but 13 Jan -16 May 1999, sample 7666, QM S74864; 1 c?, 1 same details but

Camerons Scrub, below road, 27°30 , 45” S, 152°43’39” E [±500 m], 40 m a.s.l., 11 Nov. 1998-13 Jan.

1999, sample 7555, QM S74857; 1 The Knobby, 27°30 , 27” S, 152 0 35’18” E [±500 m], 240 m a.s.l.,

II Nov. 1998-13 Jan. 1999, G. Monteith, D. Cook and G. Thompson, vine scrub, pitfall, sample 7567,

QMS74858; 1 Split Yard Creek, 27°22’41” S, 152°37 , 56” E [±500 m], 150 m a.s.l., 27 Dec. 1998-13

Jan. 1999, G. Monteith, D. Cook and G. Thompson, vine scrub, dung trap, sample 7568, QM S74859; 1

c?, Boombana, Brisbane, 27°24 , 08” S, 152°47’22” E [±100 m], 440 m a.s.l., 7 Nov. 2003, R. Raven, B.

Baehr and O. Seeman, day hand collection, ex S.C. 51695, QM S74874.

WESTERN AUSTRALIA: 1 1 §, Curtin University, Enviro nm ental Biology Building, Bentley,

[32°00 , 21” S, 115 o 53’40” E, ±200 m], 25 Apr. 2002, T. Cocket, WAM T47890; 1 S, same details

but Physics Building, 32°00 , 23” S, 115°53 , 45” E [±50 m], 10 m a.s.l., 4 Nov. 2008, M.S. Harvey, in

building, WAMT73493; 1 S, 1 Nursing Home, Hamersley Road, Subiaco [31°57’28” S, 115°49 , 25”

E, ±100 m], 30 m a.s.l., 22 Jan. 2003, K. Debnam, inside nursing home, WAM T47891; 1 §, 7 Kingston

Street, Nedlands, 31°57’56” S, 115°48’46” E [±100 m], 10 m a.s.l., 19 Apr. 2003, J.M. Waldock, found

dead on front verge near driveway, WAM T52005; 1 same details but 21 Apr. 2003, J.M. Waldock,

walking on footpath in front garden, WAM T52006; 1 same details but 19 May 2003, S. Slack-Smith,

in garden, WAM T52345; 1 same details but 28 Sep. 2007, WAM T95077; 2 same details but

6-9 Nov. 2008, WAM T94065; 2 S3, 1 juvenile, same details but 6 Nov. 2008, on path in garden after

rain, WAM T94986; 2 $ ?, 3 juveniles, same details but 8 Dec. 2008, on path in garden, WAM T94973;

1 3, 22 Windelya Road, Murdoch [32°04U4” S, 115°49’31” E, ±50 m], 30 m a.s.l., 3 Apr. 2005, O.

Mueller, WAM T63211; 1 3 , same details but 24 Apr. 2005, on verandah at night, WAM T65825; 1

5, Edward Street, Nedlands, 31°58’ S, 115°48’ E [±2km], 20 m a.s.l., 1 Mar. 2006, A. Baynes, WAM

T67503; 2 33 , Garfield Way, Greenwood, 31°49’ S, 115°48’ E [±2 km], 40 m a.s.l., 25 May 2006, T.

Struthers, garden litter, WAM T78133; 1 (?, 3 $ $, same details, WAM T112662; 1 3 , 56 Edward Street,

Nedlands [31°58’54” S, 115°48’24” E ±50 m], 20 m a.s.l., 3 May 2007, A. Baynes, WAM T82708; 1 3 ,

University of We stern Australia, Crawley, bus stop on Fairway near corner of Cooper Street, 31°58’41”

S, 115°48’56” E [±100 m], 11 Jun. 2008, J. Wojcieszek, WAM T89288; 1 3, 7 Dunkeld Street, Floreat,

31°55 , 51” S, 115°47’24” E [±100 m], 13 Dec. 2012, J. Foss, WAM T129488.

Description

(Based ma inl y on Camerons Scrub, QLD specimens.) As for the genus. Maximum male/female

midbody width ca. 3.8/4.4 mm. Colour in alcohol: “Colour brown-black, ringed tawny-pink on the

anterior (nested) end of the prozonites and on the median portion of the metazonites. Anal valves black.

Legs pale yellow” (Brolemann 1931: 295, my translation); “Bright gray-yellowish with brown posterior

margin rings on diplosegments, head mainly dark brown” (Verhoeff 1941, p. 11, my translation); base

colour in QLD and WA specimens varying from pale yellow to light brown, with narrow, dark brown

ring at rear of haplosegments, diplosegments and apodous ring; darker specimens with very narrow,

light brown rings bordering waist; narrow marginal ring of dark brown on collum, brown patches on

vertex and clypeus of head and on anal valves, antennomeres brown (darker distally), legs yellowish

(Fig. 4A-B).

Male with transverse furrows very shallow, narrow, sometimes indistinct and easily overlooked (“Pas

de sillon tranverse sur les metazonites”, Brolema nn 1931: 296; “Querfurchen sehr fein”, Verhoeff 1941:

10). No longitudinal furrows laterally on diplosegments (Fig. 4B). Sternal lamella (Fig. 2A) with sides

more or less straight, corners broadly rounded, distal margin convex. Scopulae on legs 1 to 29, i.e. not

present on last podous ring. Leg bases on rings 6 and 8 separated more widely than on other rings,

sternites a little depressed. Anterior margin of aperture with rounded-triangular, medial extension and

shorter, gently convex extension on either side (as shown in Fig. 3B for S. rubriventris).

14

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Gonocoxa ca. 1/2 telopodite length, slightly flattened anteroposteriorly and slightly concave anteriorly,

with sparse, long setae anterodistally. Cannula short, narrow, uniformly tapering towards prefemur.

Gonopod telopodite (Figs 5A-B, 6A-B, 7, 10A) reaching leg 4 bases on uncoiled specimens. Prefemur

small, rounded, densely setose posteromedially, demarcated from femorite laterally by small, narrow

notch. Remainder of telopodite clearly divided into femorite and solenomere at ca. 1/2 telopodite length.

Femorite slender, straight in posterior view, slightly convex anteriorly in lateral view, the posterior

surface very slightly concave; tipped with knob-like, rounded shoulder just lateral to lateral femorite

process, a short, wide, longitudinal groove running basally from shoulder. Fateral femorite process

a short, rounded tab directed anterodistally, close to solenomere base, extending distally a little past

femorite shoulder. Medial femorite process teardrop-shaped, the rounded base with thin, translucent

cuticle, the point directed anterodistally, terminating close to solenomere at 1/4-1/3 solenomere length.

Solenomere at base ca. 2/3 femorite width, expanded and anteroposteriorly flattened distally; a narrow,

ridge-like flange (arrow in Fig. 7D) along whole of anterior surface of solenomere, diverting medially in

short loop at ca. 1/2 solenomere length, curving sharply laterally at base to form small, rounded, anterior

extension. Solenomere divided at ca. 2/3 length by large, U-shaped indentation into medially directed

apex and distomedially directed, rounded process terminating just basal to solenomere apex; apex

divided by narrow indentation into anteroposteriorly flattened, medially directed, truncate, subapical tab

carrying opening of prostatic groove, and apical, basodistally flattened tab extending medially just past

Fig. 6. A-B. Solaenodolichopuspruvoti (Brolemann, 1931); left gonopod solenomere, posterior views.

A. Syntype, from fig. 32 in Brolemann (1931). B. QM S74841. — C. Solaenodolichopus Ipruvoth left

gonopod solenomere, posterolateral view, from fig. 3 in Jeekel (1982). Scale bar: 1 mm.

15

European Journal of Taxonomy 83: 1-36 (2014)

subapical tab. Prostatic groove (Fig. 7D) running on anterior surface of telopodite, following curve of

solenomere medial to flange, terminating at midpoint on medial margin of subapical tab.

Female with leg 2 coxa extended posteroventrally as apically rounded process (Fig. 5C-D).

Distribution

Known from open forest, rainforest and vine scrub to at least 660 m a.s.l. in the D’Aguilar Range and

adjoining areas west of Brisbane in southeast Queensland, from Mt Mee south to Ipswich, a north-

south extent of ca. 65 km (map Fig. 12). This is almost certainly the native range of the species, as very

similar, undescribed species are found elsewhere in southeast Queensland. Introduced and established

in the Perth metropolitan area in Western Australia; so far reported from Kingsley south to Kardinya,

a north-south extent of ca. 30 km. Introduced (and established?) in New Caledonia. Overlaps in range

with S. vittatus.

Taxonomic notes

The types of P. pruvoti and S. annulatus have not yet been located, and I prefer not to designate lectotypes

for these species based only on the published descriptions and illustrations of Brolemann (1931) and

Verhoeff (1941). The Brisbane and Perth specimens listed above agree closely with those descriptions

and illustrations (Figs 6A-B, 7A-B).

I suspect that the South Australian specimens described by Jeekel (1982, 2002) are an unnamed

Solaenodolichopus species, not S. pruvoti. They differ from S. pruvoti in lacking pleural keels and in

having a distinct transverse furrow on the metazonites (Jeekel 1982: 125) and a medial coxal process

A

Fig. 7. Solaenodolichopus pruvoti (Brolemann, 1931), left gonopod solenomere. — A-B. Medial views,

S. annulatus (= S. pruvoti ) syntype from fig. 2 in Verhoeff (1941) (A) and QM S74856 (B). — C-D.

Lateral (C) and anterior (D) views, QM S74857. In D, arrow points to flange on anterior surface and

dotted line marks course of prostatic groove. Scale bar: 1 mm.

16

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

on male legs 6 and 7 (Jeekel 1982: 127). The specimens described in 2002 also differ from S. pruvoti

in lacking dark transverse annulation (Jeekel 2002: 61). Both Brolemann (1931: 295, for S. pruvoti )

and Verhoeff (1941: 11, for S. annulatus ) note strongly contrasting annular rings of colour, and all

non-type adults listed above are clearly ringed (Fig. 4A-B), apart from a decoloured male in QM

S74859. Jeekel explained the difference in shape of the solenomere between S. pruvoti and the South

Australian males (Fig. 6) by saying “It should be emphasised that the distal portion of the gonopod (i.e.

the solenomerite) is quite flexible, and may be curving more or less widely in a caudal direction, and

may be crooked at the base” (Jeekel 1982: 127). It is true that the S. pruvoti solenomere can be made to

curve posteriorly by manipulation. However, in none of the preserved specimens I have so far examined

of any Solaenodolichopus species is the solenomere ‘naturally’ curved as sharply as illustrated by Jeekel

(Fig. 6C).

General notes

The collector of S. annulatus , Johan Mauritzon (1902-1950), was a lecturer in botany at Fund University

when he travelled to Australia in 1936 in company with University of Uppsala zoologist Follce Finder

(as reported in several contemporary Australian newspapers, e.g., The Adelaide Chronicle , 10 Sep.

1936: 52, http://trove.nla.gov.au/ndp/del/article/92463218. accessed 10 Oct. 2013). Mauritzon left

Fund University soon after returning from his Australian trip (H. Wittzell, in litt. 10 Oct. 2013) and his

Australian millipedes were later sent to Verhoeff from Fund by Torsten Gislen (Verhoeff 1941: 3). The

types of S. annulatus and four other species described by Verhoeff (1941) from this material have not

yet been located. The types may not have been returned to Fund, as none of the Mauritzon millipede

specimens collected in Australia are currently in the Fund University Zoological Museum (J. Elcstrom,

in litt. 13 Jan. 2014).

The reported type localities of two of Verhoeff’s five 1941 species are incorrect, and it is unclear at

which stage in the information chain from Mauritzon to Verhoeff the error appeared. Hoplatessara

( Walestessara) forceps Verhoeff, 1941 (now Cladethosoma forceps) and Hoplatria clavigera Verhoeff,

1941 were both described from Mauritzon collections in Gippsland, a district in eastern Victoria. Both

have since been collected in NSW west of Sydney, and neither has since been found in intensively

collected parts of Gippsland. However, the ‘Brisbane’ type locality for S. anmdatus is plausible, because

Verhoeff (1941: 12) gives “XII.36” as the collecting date and there are “near Brisbane” locations for

Mauritzon plant collections dated 22 Dec. 1936 in the NHRS Herbarium ( http://www.nrm.se/english/

researchandcollections/collections/databases/krvptos.8598_en.html . accessed 12 Oct. 2013) and in the

Uppsala Museum of Evolution ( http://130.238.83.220/botanik/home.php . accessed 12 Oct. 2013).

Brolemann’s 1931 paper is devoted to species collected by ‘Mme. Pruvot’, i.e., the malacologist Alice

Pruvot-Fol (1873-1972), but Brolemann explicitly named Paraulacoporuspruvoti for Pruvot-Fol’s late

husband, the zoologist Georges Pruvot (1852-1924) (Brolemann 1931: 298).

Solaenodolichopus rubriventris Verhoeff, 1928

Figs 2B, 3A-B, 8A-B, E-I

Solanodolichopus rubriventris Verhoeff, 1928: 94 (genus misspelled Solanodolichopus ), 95 (first

description, name printed in error as Solandolichopus walesius , corrected in journal Corrigenda); 114

(as Solanodolichopus rubriventris in list of species described in paper); figs 24-25 in pi. 10.

Aulacoporus rubriventris - Attems 1937: 61 (new combination), 263 (Verhoeff’s 1928 description

reworded); fig. 330 (p. 265; same as fig. 24 in Verhoeff 1928). —Jeekel 1968: 18, 29; 1981: 49.

Parwalesoma rubriventris - Jeekel 2000: 41 (new combination), 43. —Nguyen & Sierwald 2013: 1158.

17

European Journal of Taxonomy 83: 1-36 (2014)

Fig. 8. — A-B, E-I. Solaenodolichopus rubriventris Verhoeff, 1928. Lateral (A) and posterior (B)

views of gonopods in situ; left gonopod solenomere in lateral (E), anterior (F), medial (G), medial

and slightly posterior (H) and posterior (I) views. A, B, E, F, G, I from QM S74690, Ft from fig. 24 in

Verhoeff (1928). Dotted line in F marks course of prostatic groove. — C-D. S. teres (Verhoeff, 1924),

anterior views of right gonopod telopodite of lectotype, from slide mount NHRS KAS1000000005 (C)

and from fig. 7 in Verhoeff (1924) (D). Scale bars: A-B = 2 mm, C-I = 1 min

18

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Material examined

Syntypes

1 male. Upper Richmond River, NSW [27°37 S, 153°00’ E, ±10 km], Apr. 1890, R. Helms, specimen

not located; 1 $ in alcohol with original Verhoeff label, broken between rings 11 and 12, same collection

details, AM KS.76506 (formerly 47728); 1 female in alcohol with original Verhoeff label, broken

between rings 7 and 8 and rings 11 and 12, same collection details, AM KS.76507 (formerly 47723); 1

female, same collection details, specimen not located.

Other material

QUEENSLAND: 4 Tamborine Mountain [27°58’ S, 153°1U E, ±5 Ion], 22 Oct. 1912, H. Hacker,

QM S74690; 1 cJ, 1 ? in fragments, [Lamington] National Park, Macpherson Range [28°15’ S, 153°08’

E, ±5 km], 15 Dec. 1926, PA. Gilbert, AM KS.94916 (formerly K55468); 1 Eagle Heights [27°54’ S,

153° 12’ E, ±5 km], M.B. Man, 23 Mar. 1955, under logs, QM S74691; 1 $, Mt Tamborine [Tamborine

Mountain] [27°58’ S, 153 0 1U E, ±5 km], pitfall 15 Jan.M Mar. 1979, J. Grimshaw, rainforest, QM

S74692.

Description

(Based on Mt Tamborine males and syntype females.) As for the genus. Maximum male/female midbody

width ca. 5.4/6.5 mm. Colour in alcohol: “Back brownish black, without rings, only the abdomen passing

into a wine red colour” (Verhoeff 1928: 94); the least faded of the specimens examined are more or less

uniformly red-brown, darker dorsally, with a pale waist and yellow legs.

Male with with distinct transverse furrow on metazonites, stopping a little above level of ozopore. No

longitudinal furrows laterally on diplosegments. Sternal lamella (Fig. 2B) with straight sides, corners

broadly rounded, distal margin a flat inverted V. Scopulae on legs 1 to 29, i.e., not present on last podous

ring. Leg bases on ring 6 separated a little more than on ring 5; leg bases with normal separation on ring

8. Anterior margin of aperture with rounded-triangular, medial extension and shorter, gently convex

extension on either side (Fig. 3B).

Gonocoxa ca. 2/3 telopodite length, slightly flattened anteroposteriorly and slightly concave anteriorly,

with sparse, long setae anterodistally. Cannula short, narrow, uniformly tapering towards prefemur.

Gonopod telopodite (Fig. 8A-B, E-I) reaching leg 6 bases when retracted. Prefemur small, rounded,

densely setose posteromedially, demarcated from femorite laterally by small, narrow notch. Remainder

of telopodite more or less straight, slender, clearly divided into femorite and solenomere at just over

1/2 telopodite length. Femorite straight, slightly flattened anteroposteriorly, parallel-sided in posterior

view, apically thickened posteriorly as inverted triangle. Lateral femorite process small, pointed, tooth¬

like, directed distally, ca. 1/5 solenomere length. Medial femorite process teardrop-shaped, arising

near lateral process, the base expanded as thin, translucent cuticle, curving anterodistally and closely

pressed to triangular apical thickening of femorite and to solenomere base, tapering to point just

anterior to solenomere at ca. 1/2 solenomere length. Solenomere arising from anterior side of femorite;

slightly flattened anteroposteriorly with narrow, ridge-like thickening posteromedially; anteriorly

with narrow flange of cuticle arising basomedially and produced basomedially as blunt point, and at

ca. 1/3 solenomere length as short, distally acuminate process; thin, spine-like process arising at ca.

1/2 solenomere length from posteromedial ridge-like thickening, directed posterodistally and a little

medially; ridge-like thickening extended slightly at ca. 1/4 solenomere length and bearing several fine,

tooth-like projections. Solenomere expanded from ca. 3/4 length, anteroposteriorly flattened, the apex

rounded laterally, medially extended as lamellar, posteromedially directed triangle, the apical margin of

triangle expanded as narrow, basodistally flattened shelf tapering towards medial apex of triangle, and

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European Journal of Taxonomy 83: 1-36 (2014)

finely dentate medially. Prostatic groove (Fig. 8F) running just lateral to flange on anterior surface of

solenomere, terminating at apex of triangular apical tab.

Female without process on leg 2 coxa.

Distribution

Known from the forested mountains behind the Gold Coast in southeast Queensland south across the

Border Ranges to the Richmond River catchment in NSW, a north-south extent of at least 60 km (map

Fig. 12). Overlaps in range with S. vittatus.

Taxonomic notes

Verhoeff (1928: 96) wrote that he examined one male and three females, but I have not located either the

male or a third female. With the male missing, I am reluctant to designate one of the undescribed females

as lectotype. My description of male S. rubriventris is based on Verhoeff’s published notes and on

the specimens listed above from Tamborine Mountain, ca. 80 km from the roughly approximated type

locality (see below). The 1979 Tamborine Mountain male was identified as ‘ Atropisoma rubriventris ’ by

P.M. Johns during a 1987 visit to QM.

General notes

S. rubriventris , S. sulcatus and S. walesius were among millipede specimens supplied to Verhoeff by

Charles Anderson (Verhoeff 1928: 79), Director of the Australian Museum in Sydney from 1921 to

1940. Verhoeff s German manuscript on the material included descriptions of 21 new millipede species

and subspecies. It was translated into English by Anderson for the Records of the Australian Museum

and the draft translation was sent to Verhoeff for approval (Verhoeff 1928: 79). Some typographical

errors in nomenclature escaped proofreading (see synonymy above) and some morphological terms were

mistranslated. For example, Anderson consistently translated hinter in the sense of‘behind, posterior’,

even when Verhoeff used hinter to mean ‘beyond, distal’ (as in English ‘hinterland’).

Although Richard Helms (1842-1914) had been working as a collector for the Australian Museum

in Sydney in 1888 and 1889, he had gone to the Richmond River in 1890 “in the interest of a private

syndicate” (Hedley 1915: 13). I have not been able to find a report by Helms on the Upper Richmond

River trip. According to local historian Margaret Henderson (in litt., 9 Jan. 2006), “The ‘Upper Richmond

River’ usually refers to the area from Casino to the source in the ranges”. I place the collections very

approximately at Kyogle, about halfway between Casino and the Border Ranges and ca. 30 km upstream

from Casino.

Solaenodolichopus sulcatus (Verhoeff, 1928)

Figs 2C, 9

Paraulacoporus sulcatus Verhoeff, 1928: 93 (first description); fig. 11 in pi. 7.

Solaenodolichopus sulcatus - Jeekel 2000: 41 (new combination), 42, 43 (redescription from new

material); figs 3-5 (p. 48) (see taxonomic notes, below). —Nguyen & Sierwald 2013: 1160.

Paraulacoporus sulcatus - Verhoeff 1932: fig. 969 (p. 1577).

Aulacoporus sulcatus - Attems 1937: 261 (new combination), 262 (Verhoeff’s 1928 description

reworded); fig. 326 (p. 262; same as fig. 11 in Verhoeff 1928). —Jeekel 1968: 18, 29; 1981: 49.

20

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Material examined

Lectotype (here designated)

Male in alcohol with original Verhoeff label, broken between rings 8 and 9 and rings 11 and 12, North

Dorrigo, NSW [30°16’ S, 152°4r E, ±5 km], 4 Jan. 1923, A. Musgrave, AM KS.76503 (formerly

K47706).

Paralectotype

1 male, details as for lectotype, not located (see taxonomic notes, below).

Fig. 9. Solaenodolichopus sulcatus (Verhoeff, 1928) S- A. Right lateral view of midbody rings, AM

KS.77379. B-C. Lateral (B) and posterior (C) views of gonopods in situ , lectotype, AM KS.76503.

D. Posterior and slightly medial view of right gonopod solenomere, possibly of missing paralectotype,

from fig. 11 in Verhoeff (1928). E-H. Lateral (E), anterior (F), medial (G) and posterior (H) views of

left gonopod solenomere, QM S75836. Arrows in A point to longitudinal grooves, dotted line in F marks

course of prostatic groove. Scale bars: A-C = 2 mm, D-H = 1 mm.

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European Journal of Taxonomy 83: 1-36 (2014)

Other material

NEW SOUTH WALES: 1 3 , Dorrigo [30°19’ S, 152°43’E, ±5 km], W. Heron, no date, SAM (specimen

missing; described in Jeekel 2000); 1 3, Upper Styx State Forest - Petroi Forest [30°40’S 152°08’E ±10

km], 29 Jan. 1975, P. Johns and R. Killip, Nothofagus moorei forest, QM S75836; 1 3 , Bruxner Forest

Park, Coffs Harbour [30°’41” S, 153°06’03” E, ±2 1cm], 22 Mar-13 Nov. 1980 [pitfall?], G. Monteith,

sample GM 104 A/1, QM S75837; 1 3 , Leagues Scrub Flora Reserve, Oakes State Forest, 30°36’ S,

152°32’ E [±2 1cm], 780 m, 20-21 Dec. 1998, D. Biclcel, yellow pan, rainforest, AM KS.77379.

Description

As for the genus. Maximum midbody width ca. 5.0 mm Colour in alcohol: “Body brownish black,

greyish-yellow on the abdomen, back with rather broad yellowish median band, which continues over

the pro- and meta-zonites [sic] to the telson. Legs black, brownish on the joints” (Verhoeff 1928: 93);

lectotype colour now faded, prozonites light brown, metazonites and legs yellowish, pale mid-dorsal

longitudinal band ca. 1/5 maximum ring width at midbody; more recent specimens also faded, patchy

brown with pale mid-dorsal band as in lectotype.

Male lectotype with distinct transverse furrow on metazonites, stopping a little above level of ozopore

(Fig. 9A). Distinct longitudinal furrows on diplosegments (Fig. 9A) to ring 15, rising slightly anteriorly,

opening dorsally with very slight bulges below; furrow just above ozopore on pore-bearing rings. Sternal

lamella (Fig. 2C) rounded-rectangular, distal margin very slightly convex. Scopulae on legs 1 to legs of

ring 14 (lectotype) or 15 or 16 (other males examined). Leg bases on ring 6 separated a little more than

on ring 5; leg bases with normal separation on ring 8. Anterior margin of aperture more or less straight

with small, rounded-triangular, medial extension.

(Gonopod description based on lectotype and other males.) Gonocoxa a little more than 1/2 telopodite

length, slightly flattened anteroposteriorly and slightly concave anteriorly, with sparse, long setae

anterodistally. Cannula short, narrow, uniformly tapering towards prefemur. Gonopod telopodite (Fig.

9B-H) reaching leg 6 bases when retracted. Prefemur small, rounded, densely setose posteromedially,

demarcated from femorite laterally by small, narrow notch. Remainder of telopodite more or less

straight, slender, clearly divided into femorite and solenomere at just over 1/2 telopodite length.

Femorite slightly flattened anteroposteriorly, slightly concave posteriorly in lateral view, parallel-sided

in posterior view, apically a little thickened posteriorly. Femorite processes both about 1/3 solenomere

length, tapering with rounded tips; lateral process curving a little anterolaterally, medial process curving

a little anteriorly. Solenomere ca. 2/3 femorite width at base, curving slightly anterolaterally, then

posteromedially, flattened to form trough-like hollow, concave posteromedially; basal half of anterior

margin of hollow produced mediodistally as large, flat, extension with tapering, rounded tip at ca. 3/4

solenomere height; distomedially curving solenomere apex deeply notched to form rounded posterior

point separate from anteroposteriorly flattened anterior margin of hollow, the latter produced apically

as thin, subtrapezoidal tab, its medial margin roundly emarginate. Prostatic groove (Fig. 9F) running

on anterior surface of telopodite, following curve of solenomere to basal corner of subapical, anterior

margin tab, bending distally at right angle and opening at distal corner of tab.

Female not yet recognised.

Distribution

Rainforest (and eucalypt forest?) in northeast NSW from the Coffs Harbour area inland towards

Armidale, an east-west extent of ca. 90 km, at elevations to at least 780 m a.s.l. (map Fig. 12). Overlaps

in range with S. walesius.

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MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Taxonomic notes

Verhoeff (1928: 94) reported that he examined two males, but I have not located a second specimen.

The lectotype vial contains a label with ‘LECTOPARATYPE / S PM. Johns / 19.xii.67’, suggesting

that Johns may have found another male or slide mount of male parts and labelled it ‘lectotype’; this

lectotypification has not been published.

In fig. 11 of Verhoeff (1928), reproduced here as Fig. 9D, the rounded posterior process at the solenomere

apex is shown as apically notched. This appears to be an error, as the process is not notched at its tip in

the lectotype or any of the other three available S. sulcatus males. Jeekel (2000) described a fourth male,

from Dorrigo (NSW); his gonopod illustration (Jeekel’s fig. 5: 48) also shows the posterior process

without a notch. The Dorrigo male was borrowed from SAM but is currently missing from the Museum

collection (K. Sparks, in litt., 2 Jan. 2014).

General notes

The collector of the types Anthony Musgrave (1895-1959) worked as an entomologist at the Australian

Museum from 1920 until his death ( http://adb. anu.edu. au/biographv/musgrave-anthonv-7716 : accessed

30 Dec. 2013). He specialised in the taxonomy of arachnids, Diptera and Hemiptera, and is best known

as the compiler of a monumental bibliography of Australian entomology (Musgrave 1932). Musgrave’s

1923 North Dorrigo trip also yielded the types of S. walesius Verhoeff, 1928 (see below).

See also the general notes for S. rubriventris (above).

Solaenodolichopus teres (Verhoeff, 1924)

Figs ID, 2D, 3D, 8C-D

Antichiropus (Solanodolichopus) teres Verhoeff, 1924: 12 (as Antichiropus teres), 20 (as Solanodoli-

chopus teres in key), 21 (as Antichiropus (Solanodolichopus) teres, first description); figs 7-8 in pi. 1.

Aulacoporus teres Attems 1937: 261 (new combination), 263 (Verhoeff’s 1924 description reworded);

fig. 327 (p. 263; same as fig. 7 in Verhoeff 1924).

Solaenodolichopus teres- Verhoeff 1928: 94 (genus misspelled Solanodolichopus ; new combination for

teres , vittatus and vittatus dorsalis , referred to by Verhoeff as “the three forms previously described by

me”). —Jeekel 2000: 40. —Nguyen & Sierwald 2013: 1159.

Aulacoporus teres- Attems 1937: 261 (new combination), 263 (Verhoeff’s 1924 description reworded);

fig. 327 (p. 263; same as fig. 7 in Verhoeff 1924). —Jeekel 1968: 18, 29; 1971: 233 (designated as type

of Solaenodolichopus); 1981: 49.

Material examined

Lectotype (here designated)

Slide mount with original Verhoeff label containing male legpair 1, right gonopod telopodite, left gonopod

telopodite missing prefemur, left gonopod prefemur attached to the distal portion of the gonocoxa,

both legs 9 and a cuticle fragment which may be the anterior aperture margin from ring 7, NHRS-

KAS1000000005; the rest of the male body missing legpair 1, ring 7 and parts of some legs, in alcohol

but dried out in the past, somewhat distorted, ZSM A20052188; Colosseum, QLD [24°26’ S, 151°35’

E, ±5 km], E. Mjoberg, Nov.-Dec. 1912 (see taxonomic notes, below). The lectotype is the one male

specimen examined by Verhoeff, i.e., the body parts on the slide mount plus the body parts in alcohol.

Paralectotype

Female, details as for lectotype, in alcohol, body broken between rings 8 and 9, NHRS-KAS1000000007.

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European Journal of Taxonomy 83: 1-36 (2014)

Other material

None.

Description

As for the genus. Maximum midbody width ca. 3.0/3.5 mm (male lectotype/female paralectotype, but

male distorted by drying out in past). Colour in alcohol: “Body gray-yellow, on each side with wide

brown longitudinal band dorsally, becoming dark brown anteriorly. Between the brown longitudinal

bands a washed-out ochre-yellow dorsal longitudinal band with a brown median stripe. Collum dark

brown, medially more or less lighter” (Verhoeff 1924: 20, my translation); male lectotype largely

colourless, female paralectotype dull yellow grading to light reddish-brown posteriorly on each ring

with no trace of longitudinal banding.

Male with transverse furrow on metazonites, stopping well above level of ozopore. No longitudinal

furrows laterally on diplosegments. Sternal lamella (Fig. 2D) rounded-rectangular, with near-vertical

sides and more or less straight distal margin slightly raised as small, rounded projection in centre,

and with wide, shallow concavity on posterior surface on either side. Aperture details uncertain due

to Verhoeffs dissection, but if slide-mounted cuticle fragment is anterior margin of aperture, then

with gently convex extensions to anterior aperture margin on either side of rounded-triangular, medial

extension (as shown in Fig. 3B for S. rubriventris). Scopulae on legs 2-29, i.e., not present on first or

last podous rings.

Gonopod telopodite reach on venter uncertain due to dissection, and not stated by Verhoeff (1924).

Gonocoxa length relative to telopodite not known; gonocoxa fragment on lectotype slide with sparse,

long setae anterodistally. Cannula short, narrow, uniformly tapering towards prefemur. Prefemur small,

rounded, densely setose posteromedially, demarcated from femorite laterally by small, narrow notch.

Femorite (Fig. 8C) ca. 40% of telopodite length, somewhat flattened anteroposteriorly, more or less

uniformly wide to ca. 1/2 its length, then bending medially at ca. 45° and expanding, the end more or

less truncate. Lateral femorite process represented by small, short, rounded ridge at femorite tip. Medial

femorite process short, pointed, “thrust into a groove on the medial side of the base of the solenomerite

and is therefore easily overlooked” (Verhoeff 1924: 21, my translation). Solenomere (Fig. 8C-D) at

base ca. 1/2 femorite tip width, directed distally and curving laterally at midlength, distally flattened

anteroposteriorly, the lateral profile gently convex; anterior surface with thin, ridge-lilce flange along

almost entire length of solenomere, extended anterolaterally as large triangle near base and smaller

triangle just below midlength. Solenomere distomedially with U-shaped indentation, the basal rim of the

U extending as short, tooth-like projection. Solenomere distal from U curving medially, subdivided by

narrow indentation into short, rod-like apical process and anteroposteriorly flattened, truncate, subapical

process with opening of prostatic groove in middle of medial margin. Prostatic groove running on

anterior surface of telopodite, following curve of solenomere on medial side of anterior surface flange.

Female paralectotype with distolateral process (Fig. 3D) on each leg 2 coxa extending ca. 1/2 coxa

diameter laterally, with truncate lateral margin about coxa diameter in length; in situ the process resting

on ring 3 ventral surface posterior to raised end of epigyne.

Distribution

So far known only from the type locality in central coastal Queensland (map Fig. 12).

Taxonomic notes

Verhoeff based his description on one male and one female. He also examined a female 20 mm long, but

questioned whether it belonged to the same species (Verhoeff 1924: 22). The small female is a stadium

24

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

7 individual broken between rings 7 and 8 and is in the NHRS vial with the paralectotype. The lectotype

slide was labelled ‘lectotype’ by RM. Johns in 1967, but Johns did not publish this lectotypification.

Verhoeff (1924) did not give a collection date for the S. teres syntypes. Mjoberg’s unpublished field

diaries show that he arrived in Colosseum in the last week of November 1912 and left during the first

week of December 1912 (A. Ferrier, in litt., 16 Dec. 2013; see also Ferrier 2006).

The structure of the gonopod telopodite is a little unclear to me due to its flat mounting on a microscope

slide, and I have not yet seen any other Solaenodolichopus specimens with gonopods perfectly matching

those of S. teres. A male (QM S74768) pitfall-trapped in 1983 at Kroombit Tops, ca. 60 km west of

Colosseum, has a similarly short and distally bent femorite and might be conspecific. However, the

much-faded colouring of the Kroombit Tops male consists of brown transverse bands at the rear of the

metazonites against a pale yellow ground, rather than the longitudinal banding seen by Verhoeff in S.

teres. The medial femorite process in the Kroombit Tops male has a teardrop-shaped base as in S. pruvoti

(for example); details of the base are unclear in the slide-mounted S. teres gonopods.

General notes

S. teres, S. vittatns Verhoeff 1924 and A vittatus dorsalis Verhoeff 1924 were among specimens collected

by the Swedish zoologist Erik Mjoberg (1882-1938) during two expeditions to Australia, 1910-1913

(Ferrier 2006). Verhoeff obtained the Mjoberg specimens from the Naturhistoriska Riksmuseet in

Stockholm, described 43 new millipede species and subspecies from the material and published his results

in the Swedish journal Arkiv for Zoologi (Verhoeff 1924). To Verhoeff s disappointment, the journal

reproduced many of his drawings at a scale too small for clarity, among them those of Aulacoporus and

Solaenodolichopus spp.: “Die Abbildungen sind leider etwas zu stark verkleinert worden, wodurch die

Deutlichkeit mancher zarterer Teile gelitten hat” (Verhoeff 1924: 142).

Solaenodolichopus vittatus (Verhoeff, 1924)

Figs 1C-E, 2E, 10B-G

Antichiropus (Solanodolichopus) vittatus Verhoeff, 1924: 22 (first description); pi. 1, figs 5, 9, 10.

Antichiropus (Solanodolichopus) vittatus dorsalis Verhoeff, 1924: 21, 22 (first description); pi. 1, fig. 6

[New synonymy.]

Solaenodolichopus vittatus - Verhoeff 1928: 94 (genus misspelled Solanodolichopus', new combination

for teres, vittatus vittatus and vittatus dorsalis, referred to by Verhoeff as ‘the three forms previously

described by me’). — Jeekel 2000: 40. —Nguyen & Sierwald, 2013: 1160.

Aulacoporus vittatus - Attems 1937: 261 (new combination), 263 (Verhoeff s description reworded);

fig. 328 (p. 263; same as fig. 6 for S. vittatus dorsalis in Verhoeff 1924). — Jeekel 1968: 18, 29; 1981:

49.

Aulacoporus vittatus dorsalis- Verhoeff 1928:94 (genus misspelled Solanodolichopus ; new combination

for teres, vittatus vittatus and vittatus dorsalis, referred to by Verhoeff as ‘the three forms previously

described by me’). —Attems 1937: 261 (new combination), 264 (Verhoeff s description reworded). —

Jeekel 1968: 18,29; 1981:49.

Solaenodolichopus vittatus dorsalis - Jeekel 2000: 40. — Nguyen & Sierwald 2013: 1160.

Material examined

S. vittatus lectotype (here designated)

Slide mount with original Verhoeff label, containing male legpair 1, right gonopod, left gonocoxa, left

gonopod telopodite and 1 leg 9, NHRS KAS1000000004; and the rest of the male body in alcohol with

original Verhoeff label, missing legpair 1, body broken between rings 3 and 4, rings 6 and 7, and rings

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European Journal of Taxonomy 83: 1-36 (2014)

8 and 9, NHRS KAS1000000006; Mt Tambourine [Tamborine Mountain], QLD [27°58’ S, 153°11 ‘E,

±5 km], E. Mjoberg, Oct. 1912 (see taxonomic notes, below). The lectotype is the one male specimen

examined by Verhoeff, i.e. the body parts on the slide mount plus the body parts in alcohol.

S. vittatus paralectotype

Female, details as for lectotype, body broken between rings 3 and 4 and rings 5 and 6, in alcohol, NHRS

KAS 1000000006.

S. vittatus dorsalis holotype

Slide mount with original Verhoeff label, containing male legpair 1, both gonopod telopodites, 1

gonocoxa and both legs 9; and the rest of the male body in alcohol with original Verhoeff label, broken

into head+collum and rings 3, 4+5, 6+7, 8, 9+10, 11-13 and 14-telson, missing legpair 1 and legs and

gonopods from ring 7, aperture damaged; Glen Lamington [Lamington Glen], QLD [28° 15’ S, 153°0E

E, +5 km], E. Mjoberg, Nov. 1912 (see taxonomic notes, below). The holotype by monotypy is the

one male specimen examined by Verhoeff, i.e. the body parts on the slide mount plus the body parts in

alcohol, both in NHRS and registered as KAS 1000000003.

Other material

QUEENSLAND: 2 33 , Lamington National Park (‘National Park’) [28°14’ S, 153°08’ E, ±5 km], Dec.

1919, H. Hacker, QM S74826; 1 3 , Tamborine Mountain (‘Mt Tamborine’) [27°58’ S, 153°11’ E, ±5

km], Oct. 1924, A. Musgrave and G. Geissmann, QM S74827; 1 3 , same locality but 18 Mar. 1955,

M.B. Wilson, in grass, QM S74764; 1 3 , 1 ?, same locality but 26 Mar. 1955, S.B. Gunn, QM S74829;

3 33 , West Burleigh [28°07’ S, 153°26’ E, ±2 km], 12 Feb. 1953, QM personnel, QM S74828; 1 3 ,

Warrie National Park [28°13’ S, 153°16’ E, ±2 km], 2 Jul. 1971, K.R. McDonald, rainforest, under

logs, ‘S30’ on metal tag, QM S74830; 1 3 , Bahrs Scrub, 27°44’59” S, 153°10’36” E [+500 m], 200 m

а. s.l., pitfall, 29 Jan.-23 May 1981, G. Monteith, GM 1078/2, QM S74765; 1 3 , Moreton Island, 27°11’

S, 153°24’ E [+10 km], 26 Apr. 1982, W. Houston, pitfall,, yellow patch, EIS, QM S5941; 1 3 , Blue

Lagoon, Moreton Island, 27°05’35” S, 153°26’27” E [+2 1cm], 21 Sep. 1982, W. Houston, EIS, pitfall,

QM S5946; 1 3 , Springbroolc - N end, 28°09’47” S, 153°15’42” E [+500 m], 540 m a.s.l., pitfalls, 15

May-30 Aug. 1997, G. Monteith, rainforest, sample 5015, QM S74836; 1 3, Perrys Knob, 27°36’11”

S, 152°36’17” E [+500 m], 200 m a.s.l., pitfall, 15 Sep-11 Nov. 1998, G. Monteith, D. Cook and G.

Thompson, vine scrub, sample 7295, QM S74843; 1 3, same details but pitfall, 11 Nov. 1998-13 Jan.

1999, sample 7564, QM S74855; 1 3 , Enoggera Reservoir site 4, 27°26’47” S, 152°55’23” E [+100

m], 110 m a.s.l., pitfall, 7 Aug.-16 Oct. 1999, G. Monteith and J. Holt, rainforest, sample 7854, QM

S74871; 1 3 , 1 ?, 3 km SE of Kalbar, 27°57’46” S, 152°38’47” E [+100 m], 120 m a.s.l., pitfall, 2 Dec.

2000-7 May 2001, C. Burwell, sample 10160, brigalow scrub, QM S74766; 1 3 , Karawatha Forest - site

б, 27°37’33” S, 153°5’24” E [+100 m], 60 m a.s.l., 30 Apr. 2003, S. Wright and E. Volschenlc, eucalypt

woodland, sample 51168, QM S74767; 1 3 , Gold Creek Reservoir site 2, 27°28’05” S, 152°52’20” E

[+100 m], 160 m a.s.l., 4 Nov. 2003, C. Burwell, dry rainforest, sifted litter berlesate, sample 51723, QM

S74873; 2 33 , Gold Creek Reservoir site 1, 27°27’53” S, 152°52’32” E [+100 m], 140 m a.s.l., pitfall,

2-30 Jan. 2004, Queensland Museum personnel, spotted gum open forest, sample 51819, QM S74875; 1

3 , 2 JJ, same details but 30 Jan.-l Mar. 2004, sample 51939, QM S74876; 2 33 , Welsby Street, New

Farm [27°27’58” S, 153°02’56” E, ±500 m], 1 Nov. 2010, M. Shaw, pitfall, QM S95213.

Description

As for the genus. Maximum male/female midbody width ca. 3.8 (mean of 24 males, range 2.9-4.7

mm)/4.8 mm (paralectotype female). Colour in alcohol: “Body reddish yellow with two broad, blackish,

lateral longitudinal bands on the dorsum, separated by wide, reddish yellow median (band). Medial

[longitudinal] stripe washed-out reddish, ventral flanks reddish-yellow” ( vittatus , Verhoeff 1924: 20, my

translation); “Trunk black, with brown legs and wide yellow reddish dorsal median band, significantly

26

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

narrower than in vittatus. Ventral flanks black” {vittatus dorsalis , Verhoeff 1924: 21, my translation);

colour in types faded and patchy, dorsalis medial longitudinal band ca. 1/3 ring width, vittatus ca. 2/3

ring width; colour and pattern variable in other specimens (Fig. 4C-I), but always with lighter colouring

dorsally and darkest colouring on upper sides.

Fig. 10. A. — Solaenodolichopus pruvoti (Brolemann, 1931), posterior view of gonopods in situ , QM

S74858. — B-G. S. vittatus (Verhoeff, 1924). B-D, posterior view of gonopods in situ ; QM S74826 (B),

QM S95213 (C), QM S74767 (D). E-G, views of slide-mounts; left (E) and right gonopod solenomere

(F) of S. vittatus lectotype, NHRS KAS1000000004, and left (top left) and right (bottom right) gonopod

solenomere (G) of S. vittatus dorsalis holotype, NHRS KAS 1000000003. Scale bars: A-D = 2 mm, E-G

= 1 mm

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European Journal of Taxonomy 83 : 1-36 ( 2014 )

Male with transverse furrows shallow, narrow, sometimes indistinct. No longitudinal furrows laterally

on diplosegments. Sternal lamella (Fig. 2E) with sides more or less straight, corners broadly rounded,

distal margin a flat inverted V. Scopulae on legs 1 to 29, i.e., not present on last podous ring. Leg bases

on rings 6 and 8 separated more widely than on other rings, sternites a little depressed. Anterior margin

of aperture with rounded-triangular, medial extension and shorter, gently convex extension on either side

(as shown in Fig. 3B for S. rubriventris).

Gonopods usually as for S. pruvoti (Fig. 10A), but with femorite in some males thinner (Fig. 10B), and

in other males shorter and bent medially near apex, and with medial femorite process larger, reaching to

ca. 1/3-1/2 of solenomere length (Fig. 10D); intermediate forms occur (Fig. 10C).

Female with leg 2 coxa extended posteroventrally as apically rounded process (as shown for S. pruvoti

in Fig. 5C-D).

Distribution

Forest and non-forest habitats in the Brisbane region, from Moreton Island ca. 130 km south to the

Border Ranges, and inland ca. 80 km from the sea (map Fig. 12). Overlaps in range with S. pruvoti and

S. rubriventris.

Taxonomic notes

The subspecies S. vittatus dorsalis was erected for a single male only differing from S. vittatus vittatus in

size, colour of ventral flanks and width of the medial dorsal band. These three characters vary substantially

in the 24 non-type males I have examined and do not appear to be correlated. I therefore regard the

subspecies dorsalis as a synonym of the nominate subspecies vittatus. The gonopod telopodites of the

types are virtually identical (Fig. 10E-G).

S. vittatus varies more across its range than other Solaenodolichopus (described and undescribed), in

gonopod form as well as in size and colour pattern. It might better be called The S. vittatus group’, and

intensive sampling and genetic analysis across its range would be a worthwhile systematics project.

It is particularly interesting that the gonopods in S. pruvoti and some S. vittatus populations are

indistinguishable, and that no colour pattern intermediates have so far been found. When redescribing S.

pruvoti as S. annulatus , Verhoeffwrote: “Stands close to vittatus Verh. (contribution 3, figs 9 and 10) but

is distinguished by different patterning and some special features of the gonopods: the rudiment of the

tibiotarsus [rounded process on basal side of U-shaped indentation] is smaller in vittatus and not curved,

the parsolenomere [apical tab] very similar but more transversely curved, the medial displacement of the

solenomere is barely detectable, overall the solenomere is less curved and less clavate, the end in vittatus

rounded” (Verhoeff 1941: 12, my translation). These small differences are largely contained within the

variation seen in both S. pruvoti and S. vittatus. Until new diagnostic features are identified, S. vittatus

and S. pruvoti are only separable on colour pattern.

Both Verhoeff and Mjoberg spelled the type locality of S. vittatus ‘Mt Tambourine’, which in the early

20th century referred to both a village and to the small basalt plateau on which the village was located;

the modern name for the plateau is Tamborine Mountain. Verhoeff (1924) did not give a collection date

for the S. vittatus types. Mjoberg’s unpublished field diaries show that he collected at ‘Mt Tambourine’

in the last two weeks of October 1912 (A. Ferrier, in litt., 16 Dec. 2013; see also Ferrier 2006). The

lectotype slide of S. vittatus was labelled ‘lectotype’ by RM. Johns in 1967, but Johns did not publish

this lectotypification.

28

MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Verhoeffs one specimen of S. vittatus dorsalis was from ‘Glen Lamingtorf. Lamington Glen was a

locality and rainfall recording station on the Lamington Plateau in the first half of the 20th century, and

Mjoberg’s unpublished field diaries show that he collected on the ‘Lamington Plateau’ in mid-November

1912 (A. Perrier, in litt., 16 Dec. 2013; see also Perrier 2006). The holotype slide was labelled ‘holotype’

by PM. Johns in 1967.

General notes

See general notes for S. teres (above).

Solaenodolichopus walesius Verhoeff, 1928

Pigs 1A-B, 2F, 3C, 11A-C, 11E-I

Soldnodolichopus walesius Verhoeff, 1928: 94 (genus misspelled Solanodolichopus ), 95 (first

description, genus misspelled Solandolichopus ), 114 (as Soldnodolichopus walesius in list of species

described in paper); pi. 10, figs 26-27.

Parwalesoma castaneum Verhoeff, 1937: 139 (first description); figs 6-7 (p. 138).

Parwalesoma castaneum - Attems 1940: 549 (Verhoeff’s description reworded). — Jeekel 1968:

19, 29. — Jeekel 1971: 231 (noted as type of Parwalesoma). — Jeekel 1981: 49. — Jeekel 2000:

41 (synonymised with Parwalesoma walesium). — Nguyen & Sierwald 2013: 1157 (as synonym of

Parwalesoma walesium).

Aulacoporus walesius - Attems 1937: 261 (new combination), 264 (Verhoeff’s 1928 description

reworded); fig. 329 (p. 264; same as fig. 26 in Verhoeff 1928). — Jeekel 1968: 18, 29; Jeekel 1981: 49.

Parwalesoma walesium - Jeekel 2000: 41 (new combination). —Nguyen & Sierwald 2013: 1157.

Material examined

S. walesius lectotype (here designated)

Male in alcohol with original Verhoeff label, broken between rings 6 and 7 and rings 7 and 8, North

Dorrigo, NSW [30°16’S 152°41’E±5km], 4 Jan. 1923, A. Musgrave, AM KS.76508 (formerly K47704).

S. walesius paralectotypes

1 male in alcohol with original Verhoeff label, broken between rings 6 and 7, ZMB 7962; 1 female in

alcohol without original Verhoeff label, broken between rings 5 and 6, AM KS.76509 (also formerly

K47704); collection details for both as for S. walesius lectotype.

P. castaneum lectotype (here designated)

Male in alcohol with original Verhoeff label, broken between rings 8 and 9, missing left antenna, ZMB

12652; Comboyne district, NSW [31°35’ S, 152°28’ E, ±10 km], date unknown, E.C. Chisholm (see

general notes, below).

P. castaneum paralectotypes

Slide mount with original Verhoeff label containing male legpairs 1 and 9 and both gonopods, ZSM

A20033565; male rings 4-6 and an isolated diplosegment, and 1 female broken between rings 8 and

9, and 10 and 11, in alcohol with new labels (male and female in separate vials), ZSM 20052246; 1

female in alcohol with original Verhoeff label, broken between rings 6 and 7 and rings 12 and 13, AM

KS76700; collection details for all three as for P. castaneum lectotype. I assume that the male rings in

ZSM A20052246 and the slide-mounted parts in ZSM A20033565 are from the same individual, i.e.,

that together they represent a single male paralectotype specimen (see taxonomic notes, below).

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European Journal of Taxonomy 83: 1-36 (2014)

Fig. 11. — A-C, E-J. Solaenodolichopus walesius Verhoeff, 1928. A-B. Posterior views of gonopods

in situ of lectotype, AM KS.76508 (A) and Parwalesoma castaneum Verhoeff, 1937 (= S. walesius)

lectotype, ZMB 12652 (B). C. Posterior view of left gonopod solenomere tip, AM KS. 16152. E-G,

I. Lateral (E), anterior (F), medial (G) and posteromedial (I) views of left gonopod solenomere, AM

KS.93755. H. Medial view of left gonopod solenomere of P. castaneum (= S. walesius ), from fig. 6

in Verhoeff (1937). J. Posterior and slightly oblique view of left gonopod solenomere of S. walesius ,

from fig. 26 in Verhoeff (1928). — D. S. pruvoti (Brolemann, 1931), posterior view of left gonopod

solenomere tip, QM S74856 (composite image at two focus levels). Dotted line in F marks course of

prostatic groove. Scale bars: 1 mm.

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MESIBOV R., Redescriptions of Solaenodolichopus millipedes

Other material

NEW SOUTH WALES: 1 S, Comboyne [31 0 35’S, 152°28’E,±10km],Aug. 1926, E.C. Chisholm, AM

KS.94102 (see taxonomic notes, below); 2 Comboyne Cave KSS-C4, 31°34’13” S, 152°23 , 38” E

[±100 m], 21 Mar. 1971, C. Carter, AM KS.96065; 2 SS, 2 § Kerewong State Forest near Lome, site

108, 31°36’ S, 152°34 , E [±2km], pitfall, 7 Nov.-lO Dec. 1978, D. Milledge, AM KS. 16152; 1 same

details, KS. 18483; 1 S, same details but site 81, 31°35’ S, 152°38’ E j'±2 km], pitfall, 26 Sep.-12 Dec.

1978, AM KS. 105104 (ex KS.96075); 1 (?, 2 § §, Boundary Creek Road, Boundary Creek State Forest,

29°56’ S, 152°33’ E [±500 m], pitfall, 4 Feb.-9 Apr. 1993, M. Gray and G. Cassis, AM KS.93755.

Description

(Based, unless otherwise indicated, on S. walesius lectotype and paralectotypes.) As for the genus.

Maximum male/female midbody width ca. 4.5/4.5 mm Colour in alcohol: “Body ringed, the metazonites

for the most part dark brown and the prozonites wine red. / The wine-red colour of the prozonites

extends on to the anterior bands of the metazonites, which are otherwise dark brown. The lower sides

and abdomen are also wine-red, the legs yellowish” (Verhoeff 1928: 94/95); colouring of types now

faded and patchy, but rings light brown, darker dorsally, lightening to yellow at the waist, darkening

towards rear of metatergite; colouring of more recent specimens as noted by Verhoeff.

Male with distinct transverse furrow on metazonites, stopping a little above level of ozopore. No

longitudinal furrows laterally on diplosegments. Sternal lamella (Fig. 2F) broadly paraboloid in outline.

Scopulae on legs 1 to 29, i.e., not present on last podous ring. Leg bases on ring 6 separated a little more

than on ring 5; leg bases with normal separation on ring 8. Anterior margin of aperture more or less

straight with small, rounded-triangular, medial extension.

Gonocoxa (of male in AM KS.93755) about 1/2 telopodite length, slightly flattened anteroposteriorly,

with sparse, long setae anterodistally. Cannula short, narrow, uniformly tapering towards prefemur.

Gonopod telopodite (Fig. 11A-C, E-I) just reaching leg 6 bases when retracted. Prefemur small, rounded,

densely setose posteromedially, demarcated from femorite laterally by small, narrow notch. Remainder

of telopodite more or less straight, slender, clearly divided into femorite and solenomere at about 2/3

telopodite length. Femorite narrowing very slightly between ends, slightly flattened anteroposteriorly.

Medial femorite process a small, triangular tab; lateral femorite process spine-like, between 1/4 and

1/3 solenomere length, arising more distally than medial process, curving slightly anteriorly towards

solenomere. Solenomere arising anterior to femorite processes, ca. 1/2-2/3 femorite width at base,

curving laterally then medially and flattening to form trough-like hollow, concave medially; basal half

of anterior margin of hollow extending distomedially and a little anteriorly as large, flat, triangular tab

reaching ca. 1/2 solenomere length, curving posteriorly, distal margin of tab with a few small teeth;

hollow at apex of solenomere facing posteromedially, the anterior margin produced as small subapical

tab with rounded notch medially on distal margin, the solenomere apex distal to the notch at right angles

to the subapical tab. Prostatic groove running on anterior surface of telopodite and lateral to large, basal

solenomere tab, following subapical tab margin and terminating at distal corner of subapical tab.

Female without process on leg 2 coxa.

Distribution

Known from eucalypt forest (and rainforest?) in northeast NSW from the ranges northwest of Coffs

Harbour to the Comboyne district, a north-south extent of ca. 180 km (map Fig. 12). Overlaps in range

with S. sulcatus.

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European Journal of Taxonomy 83: 1-36 (2014)

Taxonomic notes

The lectotype and paralectotype S. walesius males from North Dorrigo are almost identical in size and

morphological details to the lectotype P. castaneum male from the Comboyne district, 150 km to the

south; they differ slightly in the slenderness of the femorite (Fig. 11A-B). It is a little puzzling that

Verhoeff did not recognise them as the same species. His drawings of the S. walesius telopodite tip (figs

26 and 27 in Verhoeff 1928; Fig. 11J ) appear to have been done from a posterior and slightly oblique

view of the left gonopod of the lectotype, while the drawings of the P castaneum telopodite tip (figs 6

and 7 in Verhoeff 1937; Fig. 11H) are from a medial view of the left gonopod telopodite on the lectotype

slide mount. Verhoeff may have returned one of his S. walesius males to AM before he received the

P. castaneum specimens, but he may still have held the male that found its way to ZMB. If not, it is

possible that Verhoeff had no gonopod drawings of S. walesius to compare other than his published ones.

Ironically, it was a comparison of those drawings that led Jeekel (2000) to propose that P. castaneum was

a junior synonym of S. walesius.

In both S. walesius (Fig. 11C) and S. pruvoti (Fig. 1 ID) the solenomere tip ends in an L-shaped channel.

One side of the ‘L’ is the thin, anteroposteriorly flattened tab at whose medial margin the prostatic

groove opens. The other, thicker side of the ‘L’ is distal to this tab and lies in a plane at right angles to it.

In S. pruvoti , a narrow notch (arrow in Fig. 11D) separates the two sides of the ‘L’, the distal side thus

becoming a separate process, i.e., the ‘pre-apical latero-distal process’ of Jeekel (2000). In S. walesius ,

the two sides of the ‘L’ are joined at the tip by a shallow indentation (arrow in Fig. 11C). This difference

does not seem to me to be important enough to separate two genera, which is why I have synonymised

Parw ale soma with Solaenodolichopus (see above). In future papers I will document variations at the tip

of the solenomere in other Solaenodolichopus species.

Verhoeff (1928, 1937) did not state how many specimens he examined, but gave male and female

lengths for both S. walesius and P. castaneum. My lectotypifications imply that Verhoeff looked at two

males and one female of S. walesius and two males (one dissected, most parts now missing) and two

females of P. castaneum. However, the AM whole female in alcohol (KS.76509) and the ZSM male

parts and whole female in alcohol (20052246) are missing any original Verhoeff labels. I include these

among the types because I assume that the ZSM male parts are the remains of the male dissected for

the ZSM slide mount, which does have a Verhoeff label, and because the collection details label for AM

KS.76509 appears to be original. I have not made the male S. walesius in AM KS.94102 a paralectotype

because it has only a recent, printed label; it may have been deposited directly in AM by collector E.C.

Chisholm (see below). The ZSM slide of P. castaneum was labelled ‘lectotype’ by P.M. Johns in 1967,

but this lectotypification was not published.

General notes

According to contemporary newspaper items, collector Dr E.C. Chisholm was a naturalist and

Gove rnm ent Medical Officer in the Comboyne district from 1923 to 1935 ( The Port Macquarie News

and Hastings River Advocate , 4 Jun. 1927, p. 5, http://trove.nla.gov.au/ndp/del/article/ 112527391 :

The Sydney Morning Herald , 17 Jan. 1933, p. 6, http : //trove.nla.gov.au/ndp/del/article/16945944 :

The Maitland Daily Mercury , 5 Jul. 1935, p. 3, http://trove.nla.gov.au/ndp/del/article/127096313 : all

accessed 24 Dec. 2013).

Chisholm presumably collected his Comboyne millipedes and sent them directly to Verhoeff (Verhoeff

1937: 133) during that 12-year period. As he had done with descriptions of NSW species in his 1928

paper, Verhoeff used the State museum journal. Records of the Australian Museum , as an outlet for his

descriptions of P. castaneum and four other Comboyne millipede species. Verhoeff (1937) published in

German, however, perhaps to avoid any misunderstandings arising from inadequate translation.

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MESIBOV R., Redescriptions of Solaenodolichopus millipedes

See also the general notes for S. rubriventris (above).

Discussion

As circumscribed above, Solaenodolichopus is one of Australia’s most speciose millipede genera (Car

2009; Mesibov 2008), and combined genetic and morphological studies will be needed to resolve its

internal classification. Additional collecting and genetic studies will also be needed to satisfactorily

delimit the suspiciously variable S. vittatus , and to clarify its relationship with S. pruvoti, from which it

can currently only be distinguished by colour pattern.

Fig. 12. Localities for named Solaenodolichopus species as of February 2014. — A. Australia showing

location of map B (rectangle). — B. S. teres (Verhoeff, 1924) (star), S. sulcatus (Verhoeff, 1928) (crosses)

and S. walesius Verhoeff, 1928 (open triangles), other species (filled squares); rectangle shows location

of map C. — C. S. pruvoti (Brolemann, 1931) (open squares), S. rubriventris Verhoeff, 1928 (crosses)

and S. vittatus (Verhoeff, 1924) (open triangles). Maps A-C are geographic projections; approximate

north-south map extents are 900 km for B and 220 km for C.

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European Journal of Taxonomy 83: 1-36 (2014)

Phylogenetic work would also be desirable in the larger group of eastern Australian genera, evidently

closely related, to which Aulacoporus, Solaenodolichopus and Walesoma belong. The known forms

of Aulacoporus and Walesoma (described and undescribed) are separated by ca. 1300 km, and this

disjunction is unlikely to shrink with further collecting. The range of Solaenodolichopus (map Fig. 12)

will expand considerably as new species are described and documented.

Acknowledgements

For loans of, or assistance with Verhoeff’s types I am very grateful to Janet Beccaloni (Natural

History Museum, Tondon), Cathy Car (Western Australian Museum, Perth), Jason Dunlop (Museum

fur Naturkunde, Berlin), Jonas Ekstrom (Lund University, Lund), Karin Sindemark Kronestedt

(Naturhistoriska Riksmuseet, Stockholm), Graham Milledge (Australian Museum, Sydney), Kate Sparks

(South Australian Museum, Adelaide), Jorg Spelda (Zoologische Staatssammlung, Mtinchen) and

Verena Stagl (Naturhistorisches Museum, Vienna). I am especially grateful to Dr Asa Ferrier (La Trobe

University, Melbourne) for advice on Erik Mjoberg’s Queensland expedition; to Margaret Henderson

(Research Coordinator for the Richmond River Historical Society, Lismore, NSW) for advice on the

Richmond River district; to Jean-Jacques Geoffroy (Museum National d Histoire Naturelle, Paris) for

advice on H.W. Brolemann; and to Hakan Wittzell (Lund University Botanical Museum) and Jonas

Ekstrom (Lund University Zoological Museum) for advice on Johan Mauritzon and his collections in

Australia. I thank Sandrine Feig (University of Tasmania, Hobart) for assistance with SEM work. This

study was funded by the author.

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European Journal of Taxonomy 83: 1-36 (2014)

Manuscript received: 26 February 2014

Manuscript accepted: 2 April 2014

Published on: 15 May 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, London, United

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

Denmark, Copenhagen, Denmark.

36