BONN ZOOLOGICAL BULLETIN - SUPPLEMENTUM (BzB-S),

formerly “Bonner zoologische Monographien”,

Bonn zoological Bulletin, Editor-in-Chief

Fabian Herder, Zoologisches Forschungsmuseum Alexander Koenig - Leibniz-Institut fur Biodiversitat der Tiere

(ZFMK), Ichthyology Section, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228-9122-255,

fax: +49 228-9122-212; fherder@zfmk.de

Managing Editor Supplementum Series

Ralph S. Peters, ZFMK, Hymenoptera Section, same address as above, tel. +49 228-9122-290,

fax: +49 228-9122-212; r.peters@zfmk.de

The peer-reviewed series Bonn zoological Bulletin - Supplementum, formerly “Bonner zoologische Monographien”,

has existed since 1971. It is published by the Zoological Research Museum Alexander Koenig - Leibniz Institute for

Animal Biodiversity (ZFMK), Bonn. Supplements on focus topics are produced in irregular succession. The BzB-S

consists of original zoology papers too long for inclusion in our institute’s regular journal, Bonn zoological Bulletin.

Preferred manuscript topics are: systematics, taxonomy, biogeography, anatomy, and evolutionary biology. Manuscripts

should be in American English. Authors are requested to contact the managing editor prior to manuscript submittal.

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m.bollen@zfmk.de). The online version of BzB-S is available free of charge at the BzB homepage:

http://www.zoologicalbulletin.de.

Author of this issue:

Thomas Wesener

Published: Bonn zoological Bulletin - Supplementum Vol. 63, 104 pp.

Germany.

ISBN-13: 978-3-925382-67-3

ISSN: 0302-671X

Produced by Eva-Maria Levermann, Kaiserstr. 129, 53113 Bonn, Germany; emlevermann@netcologne.de.

Printed and bound by Verlag Natur & Wissenschaft, Postfach 170209, 42624 Solingen, Germany.

Cover: The Montagne de Francais giant pill-millipede, Zoosphaerium solitarium Wesener, 2009, one of Madagascar's

numerous microendemic species, occuring only on top of a single karstic hill in northern Madagascar. Photos by Jorn

Kohler, Dannstadt.

The Giant Pill-Millipedes, order Sphaerotheriida -

An annotated species catalogue

with morphological atlas and list of apomorphies

(Arthropoda: Diplopoda)

Thomas Wesener

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Contents Volume 63

Abstract 3

Introduction 3

Historical review 3

Biogeography 4

Development and life history 5

Ecology 6

Giant Pill-Millipedes as medicine 7

Predators 7

Parasites 7

Sound production 7

Genetic studies 7

Fossil record 9

Material & Methods 9

Cataloguing 9

List of acronyms and abbreviations 9

Results 11

Characters of the order Sphaerotheriida 11

Key taxonomic characters inside the order Sphaerotheriida 13

Genus and species diversity of the order 18

Catalogue 18

Order SPHAEROTHERIIDA Brandt, 1833 18

Key to families 23

Family Procyliosomatidae Wesener & VandenSpiegel, 2009 24

Procyliosoma SiIvestri, 1917 24

Family Zephroniidae Gray, 1843 26

Subfamily Zephroniinae Gray, 1832 27

Tribe Zephroniini Jeekel, 2001 27

Bothrobelum Verhoeff, 1924 27

Cryxus Leach, 1814 27

Indosphaera Attems, 1935 28

Kophosphaera Attems, 1935 28

Leptotelopus Si Ivestri, 1897 30

Zephronia Gray, 1832 30

Tribe Sphaeropoeini Broelemann, 1913 36

Sphaerobelum Verhoeff, 1924 36

Prionobelum Verhoeff, 1924 37

Leptoprotopus Silvestri, 1897 38

Sphaeropoeus Brandt, 1833 38

Tigridosphaera Jeekel, 2000 42

Subfamily Castanotheriinae Jeekel, 1974 42

Castanotherium Pocock, 1895 42

Castanotheroides Chamberlin, 1921 49

Sechelliosoma Mauries, 1980 50

Subfamily Rajasphaerinae Hoffman, 1976 50

Rajasphaera Attems, 1935 50

Family Sphaerotheriidae Brandt, 1833 51

Sphaerotherium Brandt, 1833 51

Kylindotherium Attems, 1926 63

Family Arthrosphaeridae Jeekel, 1974 63

Arthrosphaera Pocock, 1895 63

Sphaeromimus de Saussure & Zehntner, 1902 74

Microsphaerotherium Wesener & VandenSpiegel, 2007 75

Zoosphaerium Pocock, 1895 76

Family Cyliosomatidae Jeekel, 1974 88

Cyliosoma Pocock, 1895 88

Cynotelopus Jeekel, 1986 92

Species inquirenda 92

Nomina nuda 93

Acknowledgements 93

INDEX to generic names 94

INDEX to specific names 94

References 97

Bonn zoological Bulletin - Supplementum 63: 1-104 ©ZFMK

Bonn zoological Bulletin - Supplementum 63: 1-104

August 2016

The Giant Pill-Millipedes, order Sphaerotheriida -

An annotated species catalogue

with morphological atlas and list of apomorphies (Arthropoda: Diplopoda)

Thomas Wesener

Zoologisches For sc h ungsm useum Alexander Koenig, Leibniz Institute for Animal Biodiversity,

Adenauerallee 160, 53113 Bonn, Germany, Center for Taxonomy and Evolutionary Research (Section Myriapoda);

E-Mail: t.wesener@zfmk.de

Abstract. This is the first complete catalogue documenting all species, genera, and higher categories of the giant pill-millipedes,

order Sphaerotheriida. It provides a list of all 39 known potential apomorphies of members of the order that could be used as a

template for future phylogenetic studies at the ordinal level in the Diplopoda. The catalogue also includes illustrations of key

characters of the order, as well as illustrations of the intraordinal variation of the female vulva and male posterior telopods. It

further includes a key to all five Sphaerotheriida families, short descriptions for each of the 24 genera and a summary of the

state of knowledge of the ecology and development of the giant pill-millipedes. Synonymies for all 24 valid genera are listed in

a convenient tabular format. All 351 valid Sphaerotheriida species and their synonyms are listed; 14 remain nomina dubia , while

five cannot be placed in any family. For each family, genus and species, bibliographic citations are presented, including all sub¬

sequent citations in chronological order. For each species, the number and location of type depositories as well as the locality

data from the literature are provided. Bothrobelum sulcatulum (Butler, 1872) is presented as a new combination (ex Zephronia ).

Lectotypes are designated for Zoosphaerium blandum (deSaussure & Zehntner, 1902), Z. globulus & Z. imbecillum (both syn¬

onyms of Z. voeltzkowianum (deSaussure & Zehntner, 1897)), and Z. piligerum (deSaussure & Zehntner, 1897).

Key words. Species diversity; Diplopoda; millipede

INTRODUCTION

Catalogues containing a comprehensive treatment of a tax¬

onomic group at the species level are fundamental works

needed for any form of substantial systematic or evolu¬

tionary research in zoology.

Diplopoda catalogues produced in the second half of the

20th century fonn some of the most important publica¬

tions in the field of Myriapodology. The first modern cat¬

alogue was a genus catalogue (Jeekel 1971), followed by

a classification (Hoffman 1980) and a recently updated

genus list (Shelley et al. 2000). At the ordinal level, the

first steps to species catalogues were again taken by Jeekel,

who presented the first complete overview of the orders

Siphonophorida (Jeekel 2001) and Glomeridesmida

(Jeekel 2004). Complete species catalogues at the ordinal

level are also available for Polyxenida (Nguyen Duy-

Jacquemin & Geoffroy 2003) and Callipodida (Stoev et

al. 2008). Species-rich families such as the Rhinocricidae

in the order Spirobolida (Marek et al. 2003) and the Para-

doxosomatidae in the order Polydesmida (Nguyen & Sier-

wald 2013) have been catalogued; there are also region¬

al Diplopoda catalogues for North America (Hoffman

1999) and Australia (Mesibov 2014).

The order Sphaerotheriida (giant pill-millipedes) con¬

tains five families with 351 described species and is a

moderately diverse order within the class Diplopoda. The

order, together with the orders Glomeridesmida and Glom-

erida, belongs to the basal subclass Pentazonia (Blanke &

Wesener 2014).

Historical review, classification and phylogeny: The first

giant pill-millipede species to be described was Julus

ovalis by Linne (Linnaeus 1758) in his famous 10th edi¬

tion of the Systema Naturae. The first genus-name asso¬

ciated with the Sphaerotheriida, Cryxus, was based on this

species, described by Leach (1814), but then forgotten for

more than 150 years until its revival (Jeekel 1971) and lat¬

er redescription (Golovatch et al. 2012). The first ‘classi¬

fication’ of the order was undertaken by Brandt (1833,

1841); he divided the Sphaerotheriida into the genera

Sphaerotherium and Sphaeropoeus. For a long time

species were placed into genera by their geographic ori¬

gin: all Asian species were viewed as members of the fam¬

ily Sphaeropoeidae (synonym of Zephroniidae), while the

South African, Malagasy, and Australian genera were

placed in the Family Sphaerotheriidae. It was not until the

first modern classification (Jeekel 1974) that the Indian

Zephroniidae of the genus Arthrosphaera were discovered

to be closely related to their Malagasy counterparts, be¬

coming part of the Sphaerotheriidae. The first phyloge¬

netic analysis based on numerous morphological charac¬

ters (Wesener & VandenSpiegel 2009) found the

Sphaerotheriidae to be polyphyletic, with the Australian

genus Procyliosoma standing basally to the Zephroniidae

and the remaining Sphaerotheriidae, which were divided

into the Arthrosphaeridae from India and Madagascar and

the Sphaerotheriidae sensustricto from South Africa. The

Australian genera Cyliosoma and Cynotelopus were

Thomas Wesener

Giant Pill-Millipedes, Order SPHAEROTHERIIDA

ZEPHRONIIDAE

ARTHROSPHAERIDAE

Zoosphaerium

Microsphaerotherium

Sphaeromimus

Arthrosphaera

SPHAEROTHERIIDAE

Sphaerotherium

Kyiindotherium

CYLIOSOMATIDAE

Cyliosoma

Cynotelopus

Both robe f urn

Cry xus

Phonobelum

Tigridosphaera

Sphaeroheium

Indosphaera

Kophosphaera

Zephronia

Leptotelopus

Leptoprotopus

Sphaeropoeus

Castanotherium

Cast a no them ides

SecheUiosoma

Rajasphaera

PROCYLIOSOMATIDAE ___

| Procyliosoma

Fig. 1 . Current phytogeny and classification of the giant pill-millipedes, order Sphaerotheriida. Colours correspond to those of

the distribution map (Fig. 2).

placed in the new family Cyliosomatidae (Wesener

2014a). Recently published phylogenies (Fig. 1, Table 1)

are based on 100 morphological characters (Wesener &

Vanden 2009, Wesener 2014a), and DNA (Wesener et al.

2010b).

Following recent revisions of genera (e.g. Vanden-

Spiegel et al. 2002, Wesener & Sierwald 2005b, Wesen¬

er & Wagele 2008; Golovatch et al. 2012), many named

giant pill-millipede species can now be put into an exist-

Bonn zoological Bulletin - Supplementum 63: 1-104

ing genus. The recent revisions at the generic level are al¬

so the explanation for the high number of generic level

synonymies in the order: 24 synonyms are available for

the 24 valid genera (Table 2).

Despite the renewed progress in the last decades, much

work remains to be done. For example, in the SE Asian

family Zephroniidae the genera Bothrobelum VerhoefF,

1924, Sphaeropoeus Brandt, 1833, and Tigridosphaera

Jeekel, 2000 are in need of a revision. As a result of the

©ZFMK

Species Catalogue Sphaerotheriida

Table 1 . Current classification of the Sphaerotheriida. Modified after Wesener 2014.

Order Sphaerotheriida

Procyliosomatidae (10 species)

Procyliosoma (10 species, Silvestri, 1917) E Australia and New Zealand

Zephroniidae (143 species + 1 nomen dubium)

Bothrobelum (2 species, Verhoeff, 1924) Borneo

Cryxus (1 species, Leach, 1814) Vietnam

Prionobelum (7 species, Verhoeff, 1924) Vietnam, China

Tigridosphaera (4 species, Jeekel, 2000) Malaysia

Sphaerobelum (5 species, Verhoeff, 1924) Thailand, Vietnam

Indosphaera (2 species, Attems, 1935) NE India, S. Myanmar

Kophosphaera (7 species, Attems, 1935) NE India, Nepal

Zephronia (38 species, Gray, 1832) NE India, Myanmar, Cambodia, Malaysia, Thailand

Leptotelopus (1 species, Silvestri, 1897) Myanmar

Leptoprotopus (1 species, Silvestri, 1897) Sumatra

Sphaeropoeus (20 species, Brandt, 1833) Malaysia, Vietnam, Sumatra, Borneo, Java

Castanotherium (50 species, Pocock, 1895) Borneo, Butung, Java, Sumatra, Sulawesi, Philippines

Castanotheroides (3 species, Chamberlin, 1921) Borneo, Philippines

Sechelliosoma (1 species, Mauries, 1980) Seychelles

Rajasphaera (\ species, Attems, 1935) Borneo

Sphaerotheriidae (55 species + 1 nomen dubium)

Sphaerotherium (54 species, Brandt, 1833) South Africa, Zimbabwe

Kylindotherium (1 species, Attems, 1926) South Africa

Arthrosphaeridae (106 species + 11 nomina dubia)

Arthrosphaera (40 species, Pocock, 1895) S India, Sri Lanka

Sphaeromimus (10 species, de Saussure & Zehntner, 1902) SE Madagascar

Microsphaerotherium (2 species, Wesener & VandenSpiegel, 2007) Madagascar

Zoosphaerium (54 species, Pocock, 1895) Madagascar

Cyliosomatidae (20 species)

Cyliosoma (19 species, Silvestri, 1917) W & E Australia

Cynotelopus (1 species, Jeekel, 1986) W Australia

Nomina dubia (4 species)

unclear status of the genera, it is as yet not possible to pro¬

vide a key to the genus-level taxa of the Sphaerotheriida.

In addition, there is as yet no phylogenetic framework

for the species-rich genera; a subdivision of the family

Zephroniidae, containing the bulk of Sphaerotheriida

species and genera, has not yet been attempted based on

phylogenetic methods. Furthermore, despite great progress

in some genera (South Africa: VandenSpiegel et al. 2002;

Madagascar: Wesener 2009; China: Mauries 2001; Thai¬

land: Wongthamwanich et al. 2012a), many species de¬

scriptions are still problematic and do not allow a clear

species identification. Because there are numerous species

in urgent need of revision, specimens from some areas of

distribution, such as India, Australia and Indonesia, are

close to impossible to identify without extensive revision¬

ary work.

Biogeography: Sphaerotheriida show an interesting dis¬

tribution pattern (Shelley & Golovatch 2011) and occur

in disjunct areas: eastern South Africa, Madagascar, south¬

ern India and Sri Lanka, Australia and New Zealand, the

Seychelles and Southeast Asia from Nepal and western

parts of China to the Philippines and Sulawesi (Fig. 2).

Few dispersal events are traceable in the current area

of distribution, and all natural Sphaerotheriida populations

are notably absent from any Darwinian islands (=islands

of volcanic origin). Anthropogenic dispersal events are al¬

so few and probably limited to a Sri Lankan species oc-

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

Fig. 2. Distribution map of the order Sphaerotheriida. Modified after Shelley & Golovatch 2011. Yellow = Procyliosomatidae

and Cyliosomatidae; Red = Zephroniidae; Green = Arthrosphaeridae; Blue = Sphaerotheriidae.

curring in a plantation in Tanzania (Enghoff 1977), South

African giant pill-millipedes in Malawi (VandenSpiegel

et al. 2002) and potentially Mauritius (Wesener &

Wagele 2008), as well as the suspicious occurrence of a

member of the southeast Asian family Zephroniidae on the

Seychelles islands (Golovatch & Korsos 1992, Wesener

& VandenSpiegel 2009).

Development and life history: Very little is known about

the egg and larval development of the Sphaerotheriida.

Very young larval stadia have been found in a cocoon of

dirt, indicating that eggs are wrapped in such a cocoon af¬

ter oviposition (Dohle 1992). This is similar to the treat¬

ment of eggs in the related order Glomerida (Heath et al.

1974; Iatrou & Stamou 1991). While the egg-laying

process has never been observed in giant pill-millipedes,

gravid adult females of Malagasy species have been found

with several hundred eggs inside. A few thousand eggs

were found in the gigantic females of Zoosphaerium (We¬

sener & Sierwald 2005a; personal observation). Indian

Arthrosphaera deposit 400-800 eggs (Ambarish & Srid-

har 2016). While variation at the generic or even specif¬

ic levels might occur, the recorded egg numbers in giant

pill-millipedes are higher by a factor of 10 to 100 than in

the related order Glomerida, where 12-100 eggs are de¬

posited (Heath et al. 1974). After hatching the develop-

Bonn zoological Bulletin - Supplementum 63: 1-104

ment is hemianamorphotic. First, the larvae add segments

and leg pairs after each moult, then, after reaching their

full number of segments (12) and leg pairs (21), the small

animals grow until they reach their final size and contin¬

ue to moult without any further change in their size. Af¬

ter reaching sexual maturity, both sexes continue to moult,

and, at least in some species, also to grow. The full lar¬

val development is only fragmentarily known (Enghoff et

al. 1993), e.g., the first larval stadium has never been ob¬

served. Small, immature males possess telopods that show

special characteristics of their immaturity (see Wesener &

Sierwald 2005a). Because true larvae (juvenile specimens

lacking the complete number of segments) are so rarely

encountered (personal observation), their development

most likely occurs rapidly with little time passing between

moults. Adult specimens seem to moult only once a year

(Wongthamwanich et al. 2012b; Ambarish & Sridhar

2014c; personal observation), and appear to rest for sev¬

eral weeks during the process. The moulting seems to hap¬

pen often during a time of drought, e.g. the southern hemi¬

sphere winter, June-August. The lifespan of the

Sphaerotheriida is unknown. As sexually mature females

of the larger species exist in at least four or five different

size classes, the lifespan might be as high as ten years or

more.

©ZFMK

Species Catalogue Sphaerotheriida

Ecology: Ecology and behaviour of the Sphaerotheriida

are, with a few notable exceptions, especially from India,

quite unknown. Giant pill-millipedes are mainly found in

humid forests, but they also occur in dry areas on Mada¬

gascar (the spiny forest ecosystem, e.g., Wesener 2009),

and South Africa (Schubart 1958), as well as in coastal

scrub in Western Australia (Main et al. 2002).

Sphaerotheriida seem to be absent from non-forested or

bushy areas. Giant pill-millipedes are important consumers

of leaf-litter and are known to enrich the soil of planta¬

tions in India (e.g., Ashwini & Sridhar 2002,2005,2006).

Aside from the Indian Arthrosphaera, only the giant pill-

millipedes of Thailand have been the subject of basic eco¬

logical studies (Wongthamwanich et al. 2012b). Several

studies exist on the importance of giant pill-millipedes for

improving the soil quality and as ‘composters’, especial¬

ly in India (see above).

Several species in South Africa (Haacker & Fuchs 1972)

and Madagascar (Wesener & Sierwald 2005a) are adapt¬

ed to tree-climbing and can suppress the enrolment reflex

while on branches or trees.

While some species have been found in caves (Wesen¬

er 2009, Wesener et al. 2014), Sphaerotheriida is one of

the few orders of Diplopoda in which no troglomorphic

traits, e.g., loss of colouration or ocelli, are known in any

species (Golovatch 2015).

Although Sphaerotheriida are mainly known from

forests at lower or middle elevations up to 1800 m, moun-

taintop endemics are known from Madagascar (Wesener

2009) and South Africa (Schubart 1958); species have

been recorded from the Himalayas up to 2700 m (Golo¬

vatch & Martens 1996, Wesener 2015).

Two or three species of giant pill-millipedes are found in

sympatry in most parts of their range, while up to seven

different species were recorded in the same forest on

Madagascar (Wesener 2009).

Giant pill-millipedes as medicine: Some giant pill-mil¬

lipedes are utilized in South Africa for traditional medi¬

cine. The species Sphaerotherium punctulatum, locally

called ‘Ifinyezi’ is dried, burnt and then drunk to prevent

bad luck (Jones & Jones 2000). Other species of

Sphaerotherium are regularly available in South African

alternative medicine markets (Herbert et al. 2003).

Predators of giant pill millipedes: Giant pill-millipedes

are the only large-bodied millipede taxon without defen¬

sive secretions and depend entirely on their enrolling be¬

haviour for protection (Shear 2015). Little and only an¬

ecdotal evidence exists regarding potential predators of

Sphaerotheriida. South African chlamydephorid slugs are

specialized predators of giant pill-millipedes (Herbert

2000). Generalist predators occasionally prey on giant pill-

millipedes. Ants (Formicidae) are among these general¬

ist predators, giant pill-millipede remains were found

around ant nests in Madagascar (Wesener & Sierwald

2005a); while in Cambodia individually hunting ants as

well as socially hunting species prey on giant pill-milli¬

pedes (Peeters & De Greef 2015). Large birds like Mala¬

gasy Ground Rollers or Ibis species swallow giant pill-

millipedes whole (Raseroka 1975; Thorstrom & Lind

1999). Larger mammal predators like the mongoose Ga-

lidia elegans (Wesener & Sierwald 2005a), the genet

Genetta tigrina (Roberts et al. 2007) or South African ele¬

phant shrews (Linn et al. 2007) are able to break smaller

specimens with their teeth. On Madagascar, Red-bellied

and Rufous lemurs ( Eulemur fulvus rufus and E. rubriven-

ter ) feed on tree-climbing giant pill-millipedes, apparent¬

ly just biting them into sizable pieces (Overdorff 1993).

Parasites of giant pill millipedes: Giant pill-millipedes

are hosts to internal and external parasites. Internally,

cephaline gregarines (Karandikar & Rodgi 1955, Janar-

danan & Ramachandran 1983) are often found in the in¬

testine. The nematode family Thelasmotidae with the

genus Desmicola is found in different Asian genera of gi¬

ant pill-millipedes (Hugot et al. 1991). Most parasitic

records in the Sphaerotheriida deal with nematomorphs

of the genera Rhigonema and Glomerinema , which were

recorded in species from Madagascar (Waerebeke 1984,

1985a, 1985b), Asia (Waerebeke 1986), Africa (Hunt

2002), and New Zealand (Clark 1978). Externally, at least

species from India, Madagascar and South Africa are of¬

ten found with numerous, morphologically adapted par¬

asitic mites (personal observation). Fungi of the Laboul-

beniales group grow on a New Zealand species of giant

pill-millipede (Rossi & Weir 1998), while unspecified fun¬

gi grow on Zoosphaerium arboreale from Madagascar

(Wesener & Sierwald 2005a).

Sound production: Aside from their ability to roll tight¬

ly into a ball, giant pill-millipedes, possibly uniquely in

the Diplopoda, can stridulate. Stridulation has only been

studied in the family Sphaerotheriidae, where males of

Sphaerotherium produce species-specific vibrations that

apparently overcome the enrolment reflex of the female

and allow for mating to begin (Haacker 1969, 1974, We¬

sener et al. 2011). Sound production has so far not been

studied in the family Arthrosphaeridae, where both males

and females are equipped with stridulation organs (We¬

sener & VandenSpiegel 2009). A male stridulation organ

(a third different one) seems also to exist in the monotyp-

ic zephroniid genus Leptotelopus (Pococlc 1890b).

Genetic studies: Only a handful of molecular studies (sin¬

gle genes, a mitochondrial, and a transcriptome) have been

conducted involving a giant pill-millipede species (at least

as of 01.2016). The pioneering studies were those focus¬

ing on the phylogeny of the Myriapoda in general, utiliz¬

ing two (Regier & Shultz 2001) or three nuclear protein-

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

Table 2. Genus-level synonymies in the order Sphaerotheriida.

Genus Synonym

CyliosomellaVQ rhoeff, 1924

Syncyliosoma Silvestri, 1917

Indosphaera Attems, 1936

Lophozephronia Attems, 1936

Chinosphaera Attems, 1935

Tonkinobelum Verhoeff, 1924

Pantitherium Attems, 1932

Lissosphaera Attems, 1943

Borneopoeus Verhoeff, 1924

Pulusphaera Attems, 1935

Luzonosphaera Wang, 1951

Bournellum de Saussure & Zehntner, 1902

Eubournellum Attems, 1908

Natalobelum Verhoeff, 1924

Neobournellum Attems, 1908

Oligaspis Wood, 1865

Tetraconosoma Verhoeff, 1924

Trochosoma Chamberlin, 1921

Trivandrobe/um Verhoeff 1937

Odontosternum Attems, 1943

Globotherium Brolemann, 1922

Heligmasoma Chamberlin, 1921

Epicyliosoma Silvestri, 1917

Paracyliosoma Verhoeff 1928

Valid Genus

Procyliosoma Silvestri, 1917

Indosphaera Attems, 1935

Leptotelopus Silvestri, 1897

Prionobelum Verhoeff, 1924

Sphaeropoeus Brandt, 1833

Castanotherium Pocock, 1895

Castanotheroides Chamberlin, 1921

Sphaerotherium Brandt, 1833

Arthrosphaera Pocock, 1895

Zoosphaerium Pocock, 1895

Cyliosoma Pocock, 1895

Synonymy established

Jeekel 1986: 38

Jeekel 2001: 15

Jeekel 1971: 26

Mauries 2001: 139

Jeekel 1974: 46

Hoffman 1976: 120

Jeekel 2001: 30

VandenSpiegel et al. 2002: 145

Jeekel 1971: 25

VandenSpiegel et al. 2002: 145

Jeekel 2001: 7

Jeekel 1999: 9

Wesener & Wagele 2008: 38

Wesener 2014: 199

Jeekel 1986: 36

coding genes (Regier et al. 2005). Unfortunately, as ap¬

parently no voucher specimens were deposited, the iden¬

tity of the studied species remains doubtful. An addition¬

al study focusing on arthropod phylogeny sequenced the

complete 18S ribosomal DNA gene as well as H3, U2,

COI and 16S for an Australian member of the

Sphaerotheriida (Giribet et al. 2001) whose identity is not

totally clear. The first and so far only study focusing on

the phylogeny of the Sphaerotheriida utilizing the com¬

plete 18S as well as 16S and COI (Wesener et al. 2010)

failed to provide resolution of the deeper nodes of the tree.

In this study, the 18S rDNA of the Sphaerotheriida had a

length of ca. 2000 bp, with the exception of a single

species ( Sphaeromimus musicus (deSaussure & Zehntner,

1897)) that had individual extension fragments of an ad¬

ditional 400 bp. The Zephroniidae is the only family

whose 18S gene could not be amplified so far.

Only two integrative taxonomy studies have been con¬

ducted until now (01.2016) in the Sphaerotheriida, involv¬

ing a species of Sphaerobelum Verhoeff, 1924 from Thai¬

land (Wongthamwanich et al. 2012a) and the genus

Sphaeromimus from Madagascar (Wesener et al. 2014).

Genetic distances between the few sampled Sphaerotheri¬

ida species are usually 8-20%, rarely lower in recently

specified sister-taxa (4%), while intraspecific distances

vary (0-3.5%).

Aside from single gene studies, one mitochondrial

genome has been sequenced (Dong et al. 2012). While the

authors determined the species to be a member of the

Sphaerotheriidae and no voucher was deposited, the ori¬

gin of the sampled specimen from China is an indicator

that it might represent a member of the Zephroniidae. In¬

deed, a comparison of the COI fragment shows a great

similarity to Prionobelum Verhoeff, 1924, the only giant

pill-millipede genus recorded so far from China (unpub¬

lished data).

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Species Catalogue Sphaerotheriida

A single Sphaerotheriida transcriptome has been se¬

quenced for a study focusing on hemocyanins of the Myr-

iapoda (Pick et al. 2014). The analysed species was from

the genus Arthrosphaera Pocock, 1895.

Fossil record: The Sphaerotheriida probably date back to

at least the Upper Carboniferous, where fossil records of

the related order Amynilspedida are common from North

America (Hannibal & Feldman 1981) and central Europe

(Fritsch 1901a, 1901b). Insufficiently conserved fossils

first placed in the order Sphaerotheriida are also known

from the Upper Carboniferous (Hannibal & Feldman

1981), but probably also represent Amynilspedida

(Racheboeuf et al. 2004). Fossil specimens that can be un¬

ambiguously placed in the order Sphaerotheriida are un¬

known. Additionally, probably because of their large body

size, no Sphaerotheriida are known from amber (Shear &

Edgecombe 2010).

MATERIAL & METHODS

Cataloguing. The information about species and genera

presented here is mainly derived from existing faunistic

lists (Asia: Jeekel 2001, South Africa: VandenSpiegel et

al. 2002, Australia: Mesibov 2014) and the references pro¬

vided therein. For each reference, only the first page (or

rarely the first figure) in which the species name is men¬

tioned is listed. Users of this list are advised to check the

particular reference of interest for further name-uses. As

the literature on Sphaerotheriida is of limited diversity, on¬

ly the author and year are given after each specific record;

the complete reference can be found in the “References”

section. Records are annotated with ‘list’, ‘description’,

‘new locality’, etc. The information about the types (num¬

ber, sex) is mainly taken from the first description or, more

rarely, catalogues published more recently. The provided

locality data is often restricted to the data provided in the

first description; and many Sphaerotheriida species are still

only known from their type locality. The repositories come

either from the first description or from type catalogues

of natural history collections, both published and unpub¬

lished.

At the familial and generic level, all substantive works

that can be connected to the family or genus are listed (e.g.,

publications mentioning a representative of the South

African family Sphaerotheriidae from China, where only

members of the family Zephroniidae occur, are not listed

under the Sphaerotheriidae). At the species level, it was

attempted to list all articles that mention the species name.

However, for some older publications dealing with the In¬

dian genus Arthrosphaera , species determinations seem

to be dubious, so such articles were dropped.

ACRONYMS

AM Australian Museum, Sydney, Australia.

AMNH American Museum of Natural History,

New York, U.S.A.

CAS California Academy of Sciences,

San Francisco, California, U.S.A.

CUMZ Museum of Zoology,

Chulalongkom University, Bangkok, Thailand

FMNH Field Museum, Chicago, Illinois, U.S.A.

HNMB Hungarian National Museum,

Budapest, Hungary

IEPN Instituto di Entomologia Generale et Agraria

in Portici, Napoli, Italy

IRSN Institut Royal des Sciences Naturelles de

Belgique, Brussels, Belgium

MCZ Museum of Comparative Zoology, Harvard

University, Cambridge, U.S.A.

MHNG Museum d’histoire naturelle de la Ville

de Geneve, Geneva, Switzerland

MNHN Museum national d’Histoire naturelle,

Paris, France

MRAC Royal Africa Museum Tervuren, Belgium

MTD Museum fur Tierkunde, Dresden, Germany

MZLU Lund Universitets Zoologiska Museum,

Lund, Sweden

MZUF Museo Zoologico “La Specola”,

Universita di Firenze, Italy

NHML Natural History Museum, London, England

NHMW Naturhistorisches Museum Wien,

Vienna, Austria

NHRM Naturhistoriska Riksmuseet,

Stockholm, Sweden

NMSA Natal Museum, Pietermaritzburg, South Africa

SAM South Australian Museum, Adelaide, Australia

SAMC South African Museum, Cape Town,

South Africa

SMF Forschungsinstitut und Naturmuseum

Senckenberg, Frankfurt, Germany

SMNG Senckenberg Museum fur Naturkunde Gorlitz,

Gorlitz, Germany

SMNH Swedish Museum of Natural History,

Stockholm, Sweden

WAM Western Australian Museum, Perth, Australia

ZFMK Zoologisches ForschungsmuseumA. Koenig,

Bonn, Germany

ZISP Zoological Institute, Russian Academy of

Sciences, St. Petersburg, Russia

ZMA Zoological Museum, University of

Amsterdam, Amsterdam, The Netherlands

ZMB Zoologisches Museum Berlin,

Berlin, Germany

ZMUC Zoological Museum, University of

Copenhagen, Copenhagen, Denmark

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Fig. 3. Head of the Sphaerotheriida. A = Prionobelum sp., B-E = Zoosphaerium sp. A: Frontal view (modified after Mauries

2001); B: ocelli; C: antennal groove with organ of Tomosvary and aberrant ocellus (antennae removed); D: Epipharynx, frontal

view; E: detail of organ of Tomosvary. Abbreviations: aG = antennal groove; aO = aberrant ocellus; aT = anterior tooth of epiphar¬

ynx; Co = collum (tergite 1); Gn = gnathochilarium; iT = internal tooth of epipharynx; sp = row of spines; TO = organ of Tomosvary.

ZMUH Zoological Museum, University of Hamburg, ZSM Zoologische Staatssammlung,

Hamburg, Germany Munich, Germany

ZMUM Zoological Museum,

Moscow State University, Moscow, Russia

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Species Catalogue Sphaerotheriida

Fig. 4. Gnathochilarium and mandible, SEM, Zoosphaerium sp. A: gnathochilarium ventral view; B: gnathochilarium apical view

of dorsal side; C: sensory cones of rudimentary lateral palpus; D: central located pads with ‘pillows’ of sensory cones; E: senso¬

ry cones on ‘pillow’, detail; F: right mandible, mesal view. Abbreviations: C = condylus; Ca = cardines; cP = central pads (mod¬

ified central palpi); eT = external tooth; Gu = gula; Hyp = hypopharyngeal area of gnathochilarium; iP = inner palpus; iT = inter¬

nal combed tooth; LL = lamellae lingulales; LP = rudimentary lateral palpus; Me = mentum; mp = molar plate; pL = pectinate

lamellae; St = stipites.

RESULTS

Characters of the order Sphaerotheriida

Characters identifying the Sphaerotheriida as a member

of the basal diplopod subclass Pentazonia (with orders

Glomerida and Glomeridesmida)

(1) Head capsule lacking a ‘Nackenphragma’ (plesiomor-

phy)

(2) Labrum fused to clypeus (apomorphy, joined with

Helminthomorpha)

(3) Labrum with single, triangular, anteriomedian tooth

(apomorphy)

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Fig. 5. Tergite surface (B) and posterior margin of tergite underside (=endotergum, A, C&D). A: Procyliosoma sp. (Procylioso-

matidae); B: anterior margin, Cryxus ovalis (Zephroniidae); C: Cryxus ovalis (Zephroniidae); D: Sphaerotherium similare

(Sphaerotheriidae). Abbreviation: (1) = inner area.

(4) Epipharynx loosely attached to head capsule (ple-

siomorphy)

(5) Mandible consisting of single basal joint plus gnathal

lobe (plesiomorphy)

(6) Gnathal lobe with anterior condylus, allowing an ad¬

ditional articulation (apomorphy, or myriapod ple¬

siomorphy)

(7) Gnathochilarium with fused lamellae linguales (apo¬

morphy)

(8) Body with <21 tergites, <36 pairs of legs

(9) Tergites and pleurites free (plesiomorphy)

(10) Stigmatic plates ‘divided’ or not fused (plesiomor¬

phy?)

(11) Stemites completely reduced or at least rudimenta¬

ry at midbody segments (apomorphy)

(12) Spiracles present on body segments 1 and 2 (ple¬

siomorphy)

(13) Tracheae dichotomous (apomorphy?)

(14) Intestine curved, N-shaped (apomorphy)

(15) Last male leg pair modified into telopods, consist¬

ing of pincer-shaped, three- or four-jointed structures

formed by the telopodites, and inner horns formed by

the fused syncoxite (apomorphy)

(16) First larvae hatching with three pairs of legs (ple¬

siomorphy)

Unique characters of the order Sphaerotheriida

1: Body consisting of 12 tergites plus anal shield, 11 pleu¬

rites, 21 stigmatic plates and 21 (females) or 23 (males)

pair of legs. A 13th tergite is completely fused with the

anal shield, in a few species a suture is still visible.

2: Head lacking genae, wide and short, laterally with large

antennal grooves (Fig. 3A).

3: Head tentorium lacking a transverse bar despite the

presence of an incisura lateralis (Koch 2015).

4: Each ocellarium with 60-90 ocelli arranged in a large

cluster, always with a single ocellus located laterally in

antennal groove (Figs 3B, C).

5: Organ of Tomosvary a small, round, circular pit, slight¬

ly smaller than an ocellus, located laterally in antennal

groove (Figs 3C, E).

6: Antennae placed on head laterally inside large lateral

antennal grooves (Fig. 3A).

7: Antennae short to very short, each consisting of six, usu¬

ally cylindrical segments plus disc (Fig. 3A).

8: Disc usually carrying four, but species in many genera

with 15-80 (Fig. 3A), rarely up to 186 (own observa¬

tion in Sphaeropoeus Hercules Brandt, 1833), apical

cones.

9: Antennomeres 1, rarely 5 and 6, with few isolated sen-

silla basiconica (Wesener 2009).

10: Epipharynx massive, always with a larger anterior

tooth, a large inner tooth and a fringe of spines (Fig. 3D)

11: Gnathochilarium with a normal-sized gula (Fig. 4A).

12: Gnathochilarium cardines almost completely reduced

(Fig. 4A).

13: Gnathochilarium mentum fused to undivided lamel¬

lae linguales forming a large central plate (Fig. 3A).

14: Gnathochilarium stipites large, located laterally to

lamellae linguales (Fig. 4A).

15: Gnathochilarium stipites with inner palpi only (Fig.

4B), lateral palpi rudimentary, reduced to a cluster of

4-6 tiny sensory cones (Fig. 4C).

16: Median palpi on hypopharynx transfonned into cen¬

trally located pads (Fig. 4B), lacking any sensory cones

in apical half, posterior half of underside with a unique

pad of sensory cones (Figs 4D, E).

17: Maxillary nephridia opening on the endochilarium, not

on the suture between stipes and lamellae lingulaes (Ver-

hoefF 1928, Koch 2015).

18: Mandible gnathal lobe massive, with a single exter¬

nal tooth, a 3-combed (rarely 2-combed) inner tooth, 6

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Species Catalogue Sphaerotheriida

or more pectinate lamellae and a prominently developed

molar plate with an apical groove, always lacking a

membranous fringe (Fig. 4F).

19: Mandible dorsal condylus prominent and strongly de¬

veloped (Fig. 4F).

20: Defence glands absent.

21: Underside of posterior margin of tergites (‘endoter-

gunf) often with special structures to prevent intrusion

of dirt and/or parasites between tergites while not con¬

globated (Figs 5A, C, D). Opposite part of endotergum

formed by surface of anterior margin of following ter-

gite (Fig. 5B).

22: Largest tracheae of any diplopod, originating in a large,

sclerotized tracheal pouch with three unusual muscle at¬

tachments of unclear function (Verhoeff 1928).

23: Legs consisting of sixpodomeres (coxa, prefemur, fe¬

mur, postfemur, tibia, tarsus), long and massive, the

largest of any diplopod, knife-shaped, strongly com¬

pressed laterally (Wesener 2009, 2014a).

24: Femur on posterior side always with a ridge (clean¬

ing function?).

25: Tarsi always with an apical claw and at least one spine,

always lacking a paronychium.

26: Tarsi 1 and 2 always lacking an apical spine.

27: Coxal pores absent.

28: Male leg-pairs 22 and 23 modified into anterior and

posterior telopods, respectively (Wesener & Vanden-

Spiegel 2009).

29: Both pairs of telopods with fused coxae (= syncox-

ite).

30: Anterior telopod with 3 or 4 podomeres distal to syn-

coxite, with one or several processes forming chelae or

(more often) pincers (Wesener & VandenSpiegel 2009).

31: Posterior telopods forming chelae, with 3 podomeres

distal to syncoxite, telopoditomere 3 sometimes divid¬

ed (Figs 6A-L, 7A-L).

32: Telopoditomere 2 of posterior telopods always with a

large process forming the immovable finger of chelae,

while telopoditomere 3, sometimes divided, forms the

movable finger (Figs 6A-L, 7A-L).

33: Syncoxite always with inner lobes and sharp-edged

inner horns (Figs 7A, C-F).

34: Male and female gonopore opening on posterior side

of coxa 2.

35: Male gonopore usually covered with a simple plate

(Wesener & VandenSpiegel 2009).

36: Female gonopore with a vulva consisting of two scle¬

rotized basal plates (‘bursa’) and one sclerotized apical

plate (operculum), the latter often protruding above the

coxa (Figs 8A-I).

37: Adult conglobation behaviour (=volvation). All

Sphaerotheriida species are capable of rolling into a

complete ball, with all legs, head with antennae, and first

tergite being inside (Fig. 8). All adult Sphaerotheriida

show the same modifications to roll into a ball (unlike

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the pill millipedes of the order Glomerida, where sev¬

eral different mechanisms occur, e.g. Oeyen & Wesen¬

er 2015). First tergite is dwarfed (=collum), second ter¬

gite is enlarged (thoracic shield), featuring large

grooves laterally that can encompass the tips of follow¬

ing tergites. Tergites 3-12 feature a depressed anterior

margin that can accommodate parts of preceding ter¬

gites. In many species, the underside of the tergite tips

and inner side of the anal shield carry a sclerotized ridge

that interacts with a marginal brim present on thoracic

shield, keeping volvated animals passively rolled-up.

38: Larval conglobation behaviour. Larval giant pill-mil¬

lipedes, which do not yet possess the adult number of

tergites and leg pairs yet, have morphological adapta¬

tions to volvation different from those in specimens with

the complete number of tergites. In larval specimens,

the second tergite does not possess a large lateral groove,

but a gap or schisma which encompasses the tergite tips.

Potentially, a juvenile animal would otherwise be too

short to form a closed ball, as it does not possess the

full number of tergites required to close the conglobat¬

ed ball. Additionally, in larvae the apical part of the

head, including the eyes, is often not covered by the anal

shield. Unfortunately, Sphaerotheriida larvae without the

full number of tergites are rarely observed (Dohle 1992;

Wesener, pers. obs.), which hinders a more intricate

study of their behaviour and development.

39: Spermatozoon with layer of unique palisade vesicles

(Jamieson 1997).

Taxonomic characters inside the order Sphaerotheriida

Generally, all species of the order look very similar: all

351 possess the same basic structure of the mouthparts and

head, all have the same mode of rolling-up, and all pos¬

sess the same number of tergites, pleurites, stigma-carry¬

ing plates, and leg pairs. With the exception of Bothro-

belum (whose tergites have depressions or grooves), all

genera and families of giant pill-millipedes look identi¬

cal to one another when rolled-up (Fig. 9). The few aber¬

rant forms with strongly modified mandibular and leg

joints include the monotypic genera Leptotelopus and Lep-

toprotopus. Future studies will show whether these mod¬

ifications are species-level apomorphies or merely indi¬

vidual variations.

Regarding the size, gigantic species reaching the length

of 95 mm or a diameter of 50-60 mm exist on Madagas¬

car. The largest species in other areas of distribution reach

the maximum size comparable to a ping-pong ball (Fig.

9). The smallest species are the size of a pea, while the

average size reached by the majority of species in all fam¬

ilies and genera is in between a pea and a ping-pong ball

(Fig. 9). The only exception, again from Madagascar, is

Microsphaerotherium , whose species are considerably

©ZFMK

Thomas Wesener

Fig. 6. Posterior telopods I: Families Procyliosomatidae and Zephroniidae. A: Procyliosoma sp. (after Holloway 1956); B: Zephro-

nia ovalis (after Golovatch et al. 2012); C: Kophosphaera sp. (after Attems 1936); D: Prionobelum sp. (after Mauries 2001); E:

Tigridosphaera sp. (after Jeekel 1951); F: Cryxus ovalis (after Golovatch et al. 2012); G: Sphaerobelum clavigerum (after Verho-

eff 1924); H: Rajasphaera montana (after Attems 1935); I: Sechelliosoma forcipatum (after Mauries 1980); J: Castanotherium

pellitum (after Attems 1932); K: Sphaeropoeus lugubris (after Attems 1932); L: Indosphaera feae (after Attems 1936).

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Species Catalogue Sphaerotheriida

Fig. 7. Posterior telopods II: Families Sphaerotheriidae, Arthrosphaeridae, Cyliosomatidae. A: Sphaerotherium weberii (after

Schubart 1958); B: Sphaerotherium steppense (after Schubart 1958); C: Arthrosphaera magna (after Attems 1936); D: Sphaeromimus

musicus (after Wesener & Sierwald 2005b); E: Zoosphaerium priapus (after Wesener & Wagele 2008); F: Zoosphaerium

voeltzkowianum (after Wesener & Wagele 2008); G: Zoosphaerium pseudoplatylabum (after Wesener 2009); H: Zoosphaerium

neptunus (after Wesener & Wagele 2008); I: Zoosphaerium piligerum (after Wesener & Wagele 2008); J: Zoosphaerium blandum

(after Wesener & Wagele 2008); K: Cyliosoma sp. (after Silvestri 1917); L: Cynotelopus notabilis (after Jeekel 1986).

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Fig. 8. Vulvae of different Sphaerotheriida species and genera. A: Procyliosoma sp., second leg pair with vulva (after Holloway

1956); B: Procyliosoma sp. (after Holloway 1956); C: Zephronia clivicola (after Attems 1936); D: Zephronia hirta (after Attems

1936); E: Sphaerotherium hansstroemi (after Schubart 1958); F: Sphaerotheriumperbrincki (after Schubart 1958); G: Cyliosoma

angulatum (after Wesener 2014); H: Arthrosphaera magna (after Attems 1936); I: Sphaeromimus musicus (after Wesener & Sier-

wald 2005b); J: Zoosphaerium arborealis (after Wesener & Sierwald 2005a). Abbreviations: Cx = coxa; Fe = femur; EP = later¬

al plate; IP = inner, mesal plate; O = operculum; Po = postfemur; Pre = prefemur; Ta = tarsus; Ti = tibia.

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Species Catalogue Sphaerotheriida

Sphaerotherium giganteum i cm

Fig. 9. Size variation in giant pill-millipedes, showing the extremes of their sizes. Largest known specimen, as well as largest

measured species from Madagascar (except Zoosphaerium ), South Africa and India. Average size of a Sphaerotheriida species, av¬

erage smallest size of species inside the different genera (giving Sphaeromimus inexpectatus as a reference), as well as smallest

known species. All specimens rolled-up.

Sphaerotheriida species described per Decade

Decade

Fig. 10. Species of giant pill-millipedes (order Sphaerotheriida) described per decade.

smaller and represent true dwarf forms of Sphaerotherii¬

da (Wesener & VandenSpiegel 2007). Body size is a dif¬

ficult taxonomic character, as species show subsequent

growth after reaching sexual maturity (Enghoff et al.

1993). Furthermore, in several species the largest males

are considerably smaller than the largest females (e.g.,

Zoosphaerium neptunus , see Wesener & Wagele 2008;

Wesener & Schiitte 2010).

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The most important taxonomic characters at the fami¬

ly, genus and species level are the endotergum (Figs

5A-D), the male anterior and posterior telopods (Figs

6A-L, 7A-L), and the female vulva, especially the oper¬

culum (Figs 8A-I). The posterior telopods, which are, un¬

like the gonopods of other millipede groups, not directly

involved in sperm transfer (Haacker 1969), are not always

species- or even genus-specific (Figs 6A-L, 7A-L). The

anterior telopods are more variable than the posterior ones,

and are often species-specific. Other characters of taxo¬

nomic importance at the species level are: setation of ter-

gites; number of sensory cones on the antenna; spination

on legs; length/width ratio of tarsus and femur; shape of

first stigma-carrying plate; number, position and length of

locking carinae on the anal shield.

Diversity of the order

The order Sphaerotheriida contains five families, one of

which is monogeneric (Fig. 1, Table 1). All but five (which

are nomina dubia) of the 351 valid giant pill-millipede

species are assigned to 24 genera. Six of the genera are

monotypic, while an additional three genera contain two

species. These nine species-poor genera are counterbal¬

anced by six quite species-rich genera containing 20 or

more species which are: Zoosphaerium (65 species),

Sphaerotherium (54), Castanotherium (50), Arthrosphaera

(40), Zephronia (38) and Sphaeropoeus (20). While it is

entirely possible that currently existing species-rich gen¬

era will be split or lumped together, the discovery of tru¬

ly new genera based on newly discovered species would

be surprising; the last, Microsphaerotherium , dates back

to 2007 (Wesener & VandenSpiegel 2007).

Almost no new species of Sphaerotheriida were de¬

scribed after the Second World War (1945) until recently

(Fig. 10). Several of the older descriptions, especially the

numerous ones published before 1900 (Fig. 10), are of¬

ten not suitable for species identification and require cum¬

bersome revisions of type material. Only after such revi¬

sions had recently been undertaken and the taxonomic

characters of the order were worked-out (VandenSpiegel

et al. 2002; Wesener & Sierwald 2005a, Wesener &

Wagele 2008), was it finally possible to describe new

species. Species from India, South Africa, Australia and

parts of Asia are still hard to determine because of the lack

of crucial revisions.

Nevertheless, my own observations of museum collec¬

tions suggest that many more species await their discov¬

ery. On Madagascar, several species are microendemic, re¬

stricted to a single hill or forest fragment (Wesener 2009);

it remains unknown if this microendemism is common just

on Madagascar or also in other distribution areas of the

Sphaerotheriida. On Madagascar more than 20 species of

the genus Zoosphaerium are still awaiting description,

with only females currently represented in natural histo¬

ry collections (Wesener 2006). While revisions of the

species-rich genera in the families Sphaerotheriidae and

Zephroniidae might lead to the discovery of several syn¬

onymies of widespread species, this will probably be more

than balanced by new discoveries. For example, no

Sphaerotheriida species is currently known from Laos

(Likhitrakarn et al. 2014), only one is known from Cam¬

bodia (Likhitrakarn et al. 2015) and only two are known

from Thailand (Wongthamwanich et al. 2012a). Neigh¬

bouring Vietnam (Enghoff et al. 2004) harbours at least

11 species in four genera.

As genetic distances between isolated populations of the

same species, at least on Madagascar, are reasonably high

in mitochondrial genes (2-5% COI, 2^1% 16S, see We¬

sener et al. 2010b; Wesener et al. 2014), potentially cryp¬

tic species might greatly increase the currently known and

estimated giant pill-millipede species diversity.

CATALOGUE

Order SPHAEROTHERTTDA

Brandt. 1833

Name adapted from: Sphaerotheria Brandt, 1833: 198.

Sphaerotherien C.L. Koch, 1847: 36; Sphaerotheriens

Humbert, 1865: 37; Sphaerotheria Latzel 1884: 125;

Sphaerotheriens de Saussure & Zehntner 1902: 2;

Sphaerotheridae Carl, 1906: 228; Carl 1912: 104;

Sphaerotheria Attems, 1914: 139; Attems 1926: 117;

Sphaerotheriida Jeekel, 1971: 23; Hoffman 1980: 62;

Jeekel 2001: 1; Shelley et al. 2000: 12; Wesener & Van¬

denSpiegel 2009: 545; Wesener et al. 2010b: 1184; We¬

sener et al. 2011: 967; Wesener 2014a: 199

-Zephroniadae Gray, 1843: 546; Zephronioidea Cook

1895: 2; Silvestri 1897c: 645

-Chorizocerata Verhoeff, 1910: 20; Verhoeff 1924:

40; Verhoeff 1928: 1377

Diagnosis: See above.

Included families (5)

Procyliosomatidae Wesener & Van den Spiegel, 2009

Zephroniidae Gray, 1843

Arthrosphaeridae Jeekel, 1974

Sphaerotheriidae Brandt, 1833

Cyliosomatidae Jeekel, 1974

Distribution: Australia & New Zealand, South Africa,

Madagascar, Seychelles, Southern India & Sri Lanka, SE

Asia from E. Nepal to the Philippines, SE China to Su¬

lawesi and Java (Fig. 2).

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Species Catalogue Sphaerotheriida

Fig. 11. Photographs of living Sphaerotheriida, families Procyliosomatidae and Zephroniidae. A: Procyliosoma tasmanicum Sil-

vestri, 1917; B: Castanotherium sp., walking; C: Castanotherium sp., volvated; D: Zephronia dawydoffi Attems, 1953, walking;

E: Zephronia siamensis Hirst, 1907; F: Zephronia sp., from Thailand; G: two different Zephronia species from Thailand; H: Zephro¬

nia sp., from Thailand. Fig. A (c) R. Mesibov; B-D (c) J.-P. Oeyen; G (c) P. Kautt.

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Thomas Wesener

Fig. 12. Photographs of living Sphaerotheriida, family Sphaerotheriidae & Arthrosphaeridae. A: Sphaerotherium giganteum Po-

rat, 1872; B: Sphaerotherium sp.; C: Sphaerotherium sp.; D-G: Different Arthrosphaera spp. Figs A-C (c) M. Hamer; D-G Shya-

mal Lakshminarayanan via WikimediaCommons.

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Species Catalogue Sphaerotheriida

Fig. 13. Photographs of living Sphaerotheriida, family Arthrosphaeridae, genera Arthrosphaera Pocock, 1895 and Sphaeromimus

de Saussure & Zehntner, 1902. A: three rolled-up Arthrosphaera brandti (Humbert, 1865); B: Sphaeromimus musicus (de Saus-

sure & Zehntner, 1897); C: Sphaeromimus splendidus Wesener & Sierwald, 2005; D: Sphaeromimus inexpectatus Wesener & Sier-

wald, 2005; E: the critically endangered Sphaeromimus lavasoa Wesener, 2014; F: Sphaeromimus andohahela Wesener, 2014. Fig.

A (c) J.-P. Oeyen.

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Thomas Wesener

Fig. 14. Photographs of living Sphaerotheriida, family Arthrosphaeridae, genera Sphaeromimus de Saussure & Zehntner, 1902

and Zoosphaerium Pocock, 1895. A: Zoosphaerium haackeri Wesener, 2009; B: Zoosphaerium sp.; C: Zoosphaerium sp.; D:

Zoosphaerium sp.; E: Zoosphaerium sp.; F: Zoosphaerium sp.; G: leftZ alluaudi (de Saussure & Zehntner, 1902), right Z. pseudob-

landum Wesener, 2009; H: numerous Z. blandum (de Saussure & Zehntner, 1897) with a single, shiny Sphaeromimus musicus (de

Saussure & Zehntner, 1897. Fig. B (c) J.-P. Oeyen; C-F (c) A. Hartig.

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Species Catalogue Sphaerotheriida

Fig. 15. Photographs of living or freshly conserved Sphaerotheriida, family Cyliosomatidae, genera Cyliosoma and Cynotelopus

Jeekel, 1986. A: Cyliosoma sarahae (Moir & Harvey, 2008); B: C. sarahae (left) and Cynotelopus notabilis Jeekel, 1986 (right),

Key to the families of the order Sphaerotheriida

1. - Anterior telopods modified into pair of chela. Inner

horns of posterior telopods wide, tips curved lateral¬

ly (Fig. 6A). Vulval operculum well-rounded, posi¬

tioned on lower plane than inner and lateral plates (Fig.

8A). Stigmatic plates free, no spine-like process pro¬

truding above stigma opening of previous plate. Legs

without a coxal process. Tergite 13 incompletely fused

with anal shield, laterally a large gap still visible. En-

dotergum with neither callosities, nor spines, nor hair;

marginal bristles not arranged in rows (Fig. 5A). Small

cuticular scales visible between marginal bristles and

posterior margin of endotergum. Monogeneric, 10

species. Eastern Australia and New Zealand.

.Family Procyliosomatidae

- Anterior telopods modified into pair of clamps. Inner

horns of posterior telopods with parallel tips. Vulval

operculum positioned on same level as inner and lat¬

eral plates (which can be fused). Stigmatic plates ei¬

ther almost fully covered by pleurites, or stigma open¬

ing of previous plate protected by a spine-like process.

Tergite 13 completely fused with anal shield, lateral¬

ly only a weak suture sometimes visible. Endotergum

marginal bristles arranged in rows. No cuticular scales

between marginal bristles and posterior margin of en¬

dotergum.2

2. - Inner (mesal) and lateral plates of vulva fused (Figs

8B, C). Vulval operculum well-rounded, base sur¬

rounding apical margin of basal plate (Fig. 8B). Stig¬

matic plates almost fully covered by pleurites, stigma

opening protected by coxal process. Inner area of en¬

dotergum lacking spines, setae rarely present (Fig. 5C).

All antennomeres oval, not cylindrical. 144 species in

15 genera. SE Asia and the Seychelles.

.Family Zephroniidae

- Vulva always with separate inner and lateral plates.

Base of operculum surrounded by basal plates. Stig¬

matic plates free, stigma opening protected by spine¬

like process of following plate. Endotergum inner area

with large, triangular spines and setae. Antennomeres

cylindrical, or well-rounded. Australia, eastern South

Africa, Madagascar, southern India.3

3. - Anterior telopods without sclerotized knobs. Inner

horns of posterior telopods very slender (Figs 7A, B).

Posterior telopod with stridulation organ consisting of

field with numerous sclerotized ridges on the second

podomere. Male gonopore covered by an undivided

plate and apically with a second, sclerotized plate. Vul¬

val operculum mesally elongated (Figs 8D, E). En¬

dotergum often with special callosities (Fig. 5D). All

antennomeres cylindrical. Sensorial cones on palpi of

gnathochilarium arranged in several clusters. 56

species in two genera. Eastern South Africa. Intro¬

duced populations in Zimbabwe, Malawi and possi¬

bly Mauritius.Family Sphaerotheriidae

- Anterior telopods with sclerotized knobs present on

process of second, as well as last podomere. Inner

horns of posterior telopods wide Posterior telopod

without stridulation organ. Male gonopore covered on¬

ly by an undivided plate. Vulval operculum often elon¬

gated, but then centrally, not laterally. Endotergum

never with callosities. Antennomeres cylindrical or

well-rounded. Sensorial cones on palpi of gnathochi¬

larium arranged in single clusters. 4

4. - First podomere of anterior telopod with harp carry¬

ing 1-6 stridulation ribs. Inner horns of posterior telo¬

pod with single tip. Fixed finger of posterior telopod

conically shaped, never strongly triangular. Podomere

3 not massive and strongly curved. Vulval operculum

broadly rounded (Fig. 8H), or with apical notch (Fig.

81). Inner plate of vulva strongly protruding (except

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Thomas Wesener

in Sphaeromimus). Female subanal plate enlarged to

washboard carrying 1-10 stridulation ribs on each side.

Male anal shield never with sclerotized special notch.

Antennomeres oval or cylindrical. 109 species in four

genera. Madagascar, S. India and Sri Lanka. Intro¬

duced population in the Usambara Mountains, Tanza¬

nia. Family Arthrosphaeridae

- First podomere of anterior telopod greatly enlarged and

covered with tiny spines. Inner horns of posterior telo¬

pod with double tip. Fixed finger of posterior telopod

basally wide, apically strongly tapering, strongly tri¬

angular and often much shorter than podomere 3 (Fig.

7K). Podomere 3 massive and strongly curved (Fig.

7K). Vulval operculum elongated into single tip,

shaped like isosceles triangle (Fig. 8F). Inner plate of

vulva not protruding (Fig. 8F). Female subanal plate

small, without ribs. Male anal shield always with scle¬

rotized special notch. Antennomeres cylindrical. 20

species in two genera. Coastal southwestern and east¬

ern Australia. Family Cyliosomatidae

Procvliosomatidae

Wesener & VandenSpiegeL 2009

Procyliosomatidae Wesener & VandenSpiegel, 2009: 561

(first description); Wesener et al. 2010b: 1186 (molecu¬

lar phylogenetic analysis); Wesener 2014a: 199 (morpho¬

logical phylogenetic analysis).

Type genus: Procyliosoma Silvestri, 1917

Monogeneric

10 species.

Family Procyliosoma Silvestri. 1917

Procyliosoma Silvestri, 1917: 75 (first description); At-

tems 1926: 120 (key); Attems 1928: 203 (key); Verhoeff

1932: 1379 (key); Holloway 1956: 432 (revision of New

Zealand species); Jeekel 1971: 27 (list); Jeekel 1974: 45

(list); Hoffman 1980: 63 (list); Jeekel 1986: 36 (list); Shel¬

ley et al. 2000: 12 (list); Mesibov 2000: 16 (comment);

Wesener & VandenSpiegel 2009: 561 (morphological phy¬

logenetic analysis, type genus of Procyliosomatidae); We¬

sener et al. 2010b: 1184 (molecular phylogenetic analy¬

sis); Wesener 2014a: 199 (morphological phylogenetic

analysis).

Synonyms

— Cyliosomella Verhoeff 1924: 43 (first description, in

comparison with Cyliosoma ), synonymized with Pro¬

cyliosoma by Jeekel 1986: 38. Attems 1926: 120 (list);

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Verhoeff 1928: 84 (discussion, comparison with Cylio¬

soma); Verhoeff 1932: 1382 (list); Jeekel 1971: 24 (list);

Jeekel 1974: 45 (list); Hoffman 1980: 63 (list); Jeekel

1986: 38 (synonymized with Procyliosoma); Shelley et

al. 2000: 12 (list).

Type species : Cyliosomella castaneum Verhoeff, 1924, by

monotypy

— Syncyliosoma Silvestri, 1917: 85 (introduced as sub¬

genus of Procyliosoma in key), synonymized with Pro¬

cyliosoma by Jeekel 1986: 38. Jeekel 1971: 29 (list);

Jeekel 1974: 45 (raised to genus in Sphaerotheriida re¬

vision); Hoffman 1980: 63 (list); Jeekel 1986: 38 (syn¬

onymized with Procyliosoma); Shelley et al. 2000: 12

(list).

Type species : Procyliosoma (Syncyliosoma) aurivilli Sil¬

vestri, 1917, by original designation

Type species. Procyliosoma leae Silvestri, 1917, by orig¬

inal designation.

Description: Anterior telopods modified into pair of chela.

Inner horns of posterior telopods wide, tips curved later¬

ally (Fig. 6A). Vulval operculum well-rounded, positioned

on lower plane than inner and lateral plates (Fig. 8A). Stig-

matic plates free, no spine-like process protruding above

stigma opening of previous plate. Legs without a coxal

process. Tergite 13 incompletely fused with anal shield,

laterally a large gap still visible. Endotergum with neither

callosities, nor spines, nor hair; marginal bristles not

arranged in rows (Fig. 5A). Small cuticular scales visible

between marginal bristles and posterior margin of endoter¬

gum.

Distribution: Australia (NSW, Qld, Tas); New Zealand

(NZ).

Included species and subspecies (12)

Procyliosoma andersoni (Verhoeff, 1928) [NSW]

Procyliosoma aurivilli Silvestri, 1917 [Qld]

Procyliosoma castaneum (Verhoeff, 1924) [Qld]

Procyliosoma delacyi delacyi (White, 1859) [NZ]

Procyliosoma delacyi striolatum (Pocock, 1895) [NZ]

Procyliosoma dorrigense (Verhoeff, 1928) [NSW]

Procyliosoma leae Silvestri, 1917 [Tas]

Procyliosoma leiosonum (Hutton, 1877) [NZ]

Procyliosoma tasmanicum Silvestri, 1917 [Tas]

Procyliosoma tuberculatum tuberculatum (Silvestri, 1917)

[NZ]

Procyliosoma tuberculatum westlandicum Holloway,

1956 [NZ]

Procyliosoma walesianum (Karsch, 1881) [NSW]

Procyliosoma andersoni (Verhoeff, 1928)

Cyliosomella andersoni Verhoeff 1928: 85 (first descrip.)

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Species Catalogue Sphaerotheriida

Cyliosomella andersoni — Jeekel 1981: 26 (list)

Procyliosoma andersoni.— Jeekel 1986: 39 (list, new com¬

bination).

Types: M and F syntypes.

Locality: Upper Richmond River.

Deposited: AM, KS76521.

Procyliosoma aurivillii Silvestri, 1917

Procyliosoma (Syncyliosoma) Aurivillii Silvestri, 1917: 76

(first description)

Procyliosoma (Syncyliosoma) aurivilli :-- Chamberlin

1920: 89 (list)

Procyliosoma aurivillii — Jeekel 1971: 29 (as type of Syn¬

cyliosoma)

Procyliosoma aurivillii — Jeekel 1974: 24 (list)

Procyliosoma aurivillii — Jeekel 1986: 36 (list).

Types: F holotype or syntypes.

Locality: Cape York, Queensland.

Deposited: NHRM (Silvestri 1917: 85).

Procyliosoma castaneum (Verhoeff, 1924)

Cyliosomella castaneum Verhoeff, 1924: 48 (first descrip¬

tion)

Cyliosomella castaneum — Verhoeff 1928: 85 (redescrip¬

tion)

Cyliosomella castaneum — Jeekel 1971: 24 (list)

Cyliosoma castanea — Jeekel 1981: 24 (list)

Procyliosoma castaneum :— Jeekel 1986: 36 (list, new

combination).

Types: M holotype.

Locality: Herberton, Queensland.

Deposited: NHRM?

Procyliosoma delacyi delacyi (White, 1859)

Zephronia ( Sphaerotherium ) De Lacyi White, 1859: 406

(first description)

Sphaerotherium de lacyi - Butler 1873: 177 (list)

Sphaerotherium De Lacyi — Preudhomme de Borre 1884:

26 (list)

Cyliosoma de lacyi — Pocock 1895: 415 (new comb.)

Cyliosoma Delacyi:— Attems 1914: 64 (list)

Procyliosoma delacyi delacyi :— Holloway 1956: 435 (new

combination, redescription)

Procyliosoma delacyi delacyi :— Shields 2008: 33 (feed¬

ing)

— Sphaerotherium novae-zealandidae Kirk, 1885: 139,

synonymized by Holloway 1956: 441

Procyliosoma Novae-zealandiae — Silvestri 1917: 84 (re¬

description).

Types: P. delacyi 1 F, S. novae-zealandidae unknown

Locality: P. delacyi'. New Zealand (Holloway 1956: 42).

Deposited: P. delacyi NHML (Holloway 1956: 42),

Sphaerotherium novae-zealandidae specimens NHML?

Procyliosoma delacyi striolatum (Pocock, 1895)

Cyliosoma striolatum Pocock, 1895: 414 (first description)

Cyliosoma striolatum — Attems 1914: 64 (list)

Procyliosoma striolatum — Silvestri 1917: 62 (new com¬

bination)

Procyliosoma Delacyi — Silvestri 1917: 82 (redescription)

NOT White, 1859

Procyliosoma delacyi striolatum — Holloway 1956: 442

(effectively listing P. striolatum as a subspecies under

P. delacyi).

Types: F holotype.

Locality: Greymouth, South Island, New Zealand (Pocock,

1895:415)

Deposited: P. striolatum , NHML 1889.11.6.1 (Holloway

1956: 443, no longer present? J. Beccaloni, pers. comm.).

Procyliosoma dorrigense (Verhoeff, 1928)

Cyliosomella andersoni dorrigense Verhoeff, 1928: 86

(first description)

Cyliosomella andersoni dorrigense :— Jeekel 1981: 26

(list)

Procyliosoma dorrigense :— Jeekel 1986: 39 (list; raised

to species, new combination).

Types: M holotype.

Locality: North Dorrigo, NSW.

Deposited: AM.

Procyliosoma leae Silvestri, 1917

Procyliosoma (Procyliosoma) Leae Silvestri, 1917: 75

(designated type of Procyliosoma , first description)

Procyliosoma leae — Chamberlin 1920: 89 (list)

Procyliosoma leae — Jeekel 1971: 27 (list)

Procyliosoma leae — Jeekel 1981: 27 (list)

Procyliosoma leae — Jeekel 1986: 36 (list)

Procyliosoma leae — Mesibov 2000: 16 (list)

Procyliosoma leae — Wesener & VandenSpiegel 2009: 558

(morphological phylogenetic analysis)

Procyliosoma leae - Wesener et al. 2010b: 1189 (mole¬

cular phylogenetic analysis).

Procyliosoma leae — Wesener 2014a: 203 (morphologi¬

cal phylogenetic analysis).

Types: M and F syntypes.

Locality: Hobart, Tasmania

Deposited: IEPN?

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Thomas Wesener

Procyliosoma leiosomum (Hutton, 1877)

Sphaerotherium leiosomum Hutton, 1877: 116 (first de¬

scription)

Sphaerotherium leiosomus — Hutton 1878: 291 (descrip¬

tion)

Sphaerotherium leisomum [sic]:— Preudhomme de Borre

1884: 27 (list)

Cyliosoma liosoma [sic]:— Pocock 1895: 415 (list, new

combination)

Cyliosoma leisomum — Attems 1914: 64 (list)

Procyliosoma leiosoma — Silvestri 1917: 79 (redescrip.)

Procyliosoma leiosomum.— Holloway 1956: 435 (re¬

description).

Types: M and F syntypes.

Locality: Dunedin, New Zealand.

Deposited: Otago Museum, not labelled as types (Hol¬

loway 1956: 435), NHML, types or Pocock’s specimens.

Procyliosoma tasmanicum Silvestri, 1917

(Fig. 11 A)

Procyliosoma (Procyliosoma) tasmanicum Silvestri, 1917:

76 (first description)

Procyliosoma tasmanicum :— Chamberlin 1920: 89 (list)

Procyliosoma tasmanicum :— Jeekel 1981: 27 (list)

Procyliosoma tasmanicum :— Jeekel 1986: 39 (list)

Procyliosoma tasmanicum :— Mesibov 2000: 16 (list).

Types: M and F syntypes.

Locality: Hobart, Tasmania.

Deposited: IEPN?

Procyliosoma tuberculatum tuberculatum Silvestri, 1917

Procyliosoma tuberculatum Silvestri, 1917: 80 (first de¬

scription)

Procyliosoma tuberculatum tuberculatum :— Holloway

1956: 436 (redescription, designation of subspecies)

Procyliosoma tuberculatum tuberculatum :— Clark 1978:

1 (parasitic Nematoda)

Procyliosoma tuberculatum :— Rossi & Weir 1998: 282

(Funghi)

Procyliosoma tuberculata [sic]:— Clair 2011: 75 (rat pre¬

dation).

Types: M and F syntypes.

Locality: Cape Maria van Diemen, New Zealand.

Deposited: IEPN?

Procyliosoma tuberculatum westlandicum Holloway,

1956

Procyliosoma tuberculatum westlandicum Holloway,

1956: 439 (first description).

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Types: 1 F holotype, 1 M, 3F paratypes.

Locality: Buller Creek, Punakaiki, Westland, New Zealand

(Holloway 1956: 439)

Deposited: Canterbury Museum, holotype (Myr. 70, Myr

70-1, Myr 70-2), allotype (Myr 1, Myr 1-1, Myr 1-2), 2

F paratypes (Myr 5, Myr 5-1, Myr 71, Myr 71-1, Myr 71-

2), Dominion Museum, 1 F paratype (8/203,9/218) (Hol¬

loway 1956: 439).

Procyliosoma walesianum (Karsch, 1881)

Sphaerotherium walesianum Karsch, 1881: 31 (first de¬

scription)

Sphaerotherium walesianum :— Preudhomme de Borre

1884: 28 (list)

Sphaerotherium walesianum :— Pocock 1893: 130 (syn-

onymized with S. angulatum ; Jeekel (1986: 36) believes

this was based on misidentification)

Sphaerotherium walesianum.— Attems 1914: 141 (list)

Sphaerotherium walesianum — Silvestri 1917: 62 (list,

c =Cyliosoma walesianum’ \ new combination)

Cyliosoma walesianum :— Jeekel 1981: 26 (list)

Procyliosoma walesianum — Jeekel 1985: 39 (list; new

combination)

Types: M holotype.

L ocal ity: Sydney, N SW.

Deposited: ZMB 370.

Family Zephroniidae Gray. 1843

Zephroniadae Gray, 1843: 546; Zephroniidae Pocock,

1894: 326 (raised to family); Jeekel 1971: 29; Jeekel 2001:

12; Wesener & VandenSpiegel 2009: 551 (morphological

phylogenetic analysis); Wesener et al. 2010: 1188 (mol¬

ecular phylogenetic analysis); Golovatch et al. 2012: 273;

Wesener 2014a: 199 (morphological phylogenetic analy¬

sis).

— Sphaeropoeidae Brolemann, 1913: 79; Attems 1914:

142; Jeekel 1971: 28; Jeekel 1974: 46; Hoffman 1976:

118; Hoffman 1980: 63; Shelley et al. 2000: 12; Mauries

2001: 140; Wesener & Sierwald 2005b.

Remark: This group was previously known under the

name Sphaeropoeidae. With the inclusion of the genus

Zephronia in the family, and a redescription of the latter

(Golovatch et al. 2012), the name Zephroniidae takes pri¬

ority. Previously, all Asian genera were included in the

family, later Arthrosphaera was split-off (Jeekel 1974).

The Australian genus Cynotelopus was accidentally

placed in the family later (Shelley et al. 2000).

©ZFMK

Species Catalogue Sphaerotheriida

Genera included (15)

Bothrobelum Verhoeff, 1924 (Borneo)

Cryxus Leach, 1814 (‘Indochina’)

Indosphaera Attems, 1935 (NE India, S. Burma)

Kophosphaera Attems, 1935 (NE India, Nepal)

Leptotelopus Silvestri, 1897 (Burma)

Zephronia Gray, 1832 (NE India, Nepal, Burma, Cambo¬

dia, Malaysia, Thailand)

Sphaerobelum Verhoeff, 1924 (Thailand, Vietnam)

Prionobehm Verhoeff, 1924 (Vietnam, China)

Leptoprotopus Silvestri, 1897 (Sumatra)

Sphaeropoeus Brandt, 1833 (Malaysia, Vietnam, Suma¬

tra, Borneo, Java)

Tigridosphaera Jeekel, 2000 (Malaysia)

Castanotherium Pocock, 1895 (Halmahera, Butung, Bor¬

neo, Java, Sumatra, Sulawesi, Philippines)

Castanotheroides Chamberlin, 1921 (Borneo, Philippines)

Sechelliosoma Mauries, 1980 (Seychelles)

Rajasphaera Attems, 1935 (Borneo)

Type genus: Zephronia Gray, 1832

143 species + 1 nomina dubia.

Remarks: Here treated differently than the other families;

subfamilial categories after Jeekel (2001) are kept in this

list of the family Zephroniidae. However, such groupings

have not been tested yet by a phylogenetic analysis and

are arbitrary at best.

Subfamily Zephroniinae Gray, 1832

Name adapted from Zephroniadae Gray, 1843: 546; Jeekel

2001 : 12

— Sphaeropoeinae; Jeekel, 1974: 46; Hoffman 1976: 118;

Hoffman 1980: 63; Mauries 2001: 140

Tribe Zephroniini Jeekel, 2001

Zephroniini Jeekel, 2001: 13 name adapted from Zephro¬

niadae Gray, 1843: 546

— Indosphaerini Hoffman, 1976: 119; Hoffman 1980: 63

— Prionobelini Hoffman, 1976: 119; Hoffman 1980: 63

— Sphaeropoeini Mauries, 2001: 140

Bothrobelum Verhoeff. 1924

Verhoeff, 1924: 58 (first description); Attems 1926: 120

(list); Attems 1928: 203 (list); Jeekel 1971: 23 (list); Jee¬

kel 1974: 46 (classification); Hoffman 1976: 120 (list);

Hoffman 1980: 64 (list); Jeekel 2001: 13 (list); Wesener

& VandenSpiegel 2009: 551 (list).

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Description: Tergites sulcate, with a U-shaped depression

and six oval pits on each side, a unique character for the

order.

Type-species: Bothrobelum rugosum Verhoeff, 1924.

Species included (2)

Bothrobelum rugosum Verhoeff, 1924

Bothrobelum sulcatulum (Butler, 1872) n. comb.

Bothrobelum rugosum Verhoeff, 1924

Bothrobelum rugosum Verhoeff, 1924: 67 (first descrip.)

Bothrobelum rugosum.— Jeekel 2001: 13 (list).

Types: F holotype.

Habitat: Indonesia: Kalimantan, Landak.

Deposited: Unknown.

Bothrobelum sulcatulum (Butler, 1872) new combination

Zephronia sulcatula Butler, 1872: 357 (first description)

Zephronia sulcatula :— Preudhomme de Borre 1884: 25

(list)

Sphaeropoeus sulcatulus :— Silvestri 1896a: 20 (list)

Zephronia sulcatula Attems:— 1914: 147 (list)

Zephronia sulcatula — Jeekel 2001: 21 (list).

Remarks: R. L. Hoffman wrote in October 2011 ‘I looked

at a number of old dry types in the British Museum, and

noticed the odd looking one named Zephronia sulcatula

Butler. A pencil note made at the time says “Each tergum

with median u-shaped depression and 6 large oval pits on

each side.”‘ According to this letter, the species is placed

in the genus Bothrobelum.

Types: Holotype of unknown sex.

Habitat: “Borneo”, Sarawak.

Deposited: NHML (online catalogue)

Crvxus Leach. 1814

Cryxus Leach, 1814: 407; Jeekel 1971: 24 (list); Jeekel

2001: 14 (list); Wesener & VandenSpiegel 2009: 551 (list);

Golovatch et al. 2012: 277 (redescription)

Description: Anterior telopods with three podomeres dis¬

tal to syncoxite, telopoditomere 3 heavily bent posterior¬

ly, forming a ‘beak’ with the process of telopoditomere 2.

Posterior telopod, telopoditomere 2 strongly elongated,

longer than telopoditomeres 3 and 4 combined. Telopodit¬

omere 4 almost rudimentary, with a 90% torsion so that

it overlaps the immovable finger. Tarsus slender, with a

single apical spine. Small-bodied, only up to 20 mm long.

©ZFMK

Thomas Wesener

Type-species: Julus ovalis Linnaeus, 1758.

Monotypic

Cryxus ovalis (Linnaeus, 1758)

Julus ovalis Linnaeus, 1758: 639 (first description)

Julus ovalis.— Linnaeus 1759: 215 (description)

Julus ovalis — Olivier 1792: 412 (description)

Glomeris ovalis — Latreille 1804: 64 (new combination)

Glomeris ovalis — Latreille 1810: 423 (list)

Cryxus ovalis Leach 1814: 407 (new combination);

Sphaerotherium ovale - Brandt, 1841a: 363 (new com¬

bination)

Sphaerotherium ovale — Brandt 1841b: 180 (list)

Sphaerotherium ovale — Gervais 1847: 85 (list)

Sphaerotherium ovale — Butler 1873: 178 (list)

Sphaerotherium ovale — Preudhomme de Borre 1884: 27

(list)

Cryxus ovalis — Jeekel 2001: 14 (list)

Cryxus ovalis — Golovatch et al. 2012: 278 (redescription)

Cryxus ovalis — Wesener 2014a: 204 (morphological phy¬

logenetic analysis)

— Julus ovatus Fabricius, 1775: 427 (first description),

synonym, under C. ovalis by Golovatch et al. 2012: 278

Julus ovatus — Fabricius 1781: 528 (list)

Julus ovatus — Fabricius 1793: 393 (list).

Remarks: Apparently forgotten between 1847 and 1971,

redescribed in 2012 (Golovatch et al. 2012).

Types: M neotype (designated in Golovatch et al. 2012:

278).

Habitat: Asia, ‘China’, Vietnam, Cat Tien NP.

Deposited: ZMUM.

Indosphueru Allans. 1935

Indosphaera Attems, 1935: 128 (first description); Attems

1943: 60 (list); Rajulu 1970: 128 (list); Jeekel 1971: 25

(list); Jeekel 1974: 46 (classification); Hoffman 1976: 119

(list); Hoffman 1980: 3 (list); Jeekel 2001 (list): 15; Mau-

ries2001: 141 (list); Wesener & VandenSpiegel 2009: 551

(list).

Synonym

— Indosphaera Attems, 1936: 152 (first description);

Jeekel, 1971:25 (list).

Type species : Indosphaera curiosa Attems, 1936 [syn-

onymized by Jeekel 2001: 15 with Indosphaera feae\.

Description: Coxae 2 of female completely fused, a

unique character in the order. Anterior telopod with four

podomeres distal to syncoxite. Telopoditomeres 3 and 4

each almost as long as telopoditomere 1. Process of sec¬

ond telopoditomere of anterior telopods located posteri¬

orly. Posterior telopod with four telopoditomeres, immov¬

able finger with two membranous ledges. Operculum of

vulva very narrow. Tarsi with a single apical spine. Ter-

gites hairy. 35—40 mm long.

Type-species: Indosphaera feae Attems.

Species included (2)

Indosphaera feae Attems, 1935 (India, Assam)

Indosphaera fortis Attems, 1936 (Burma, Shan States)

Indosphaera feae Attems, 1935

Indosphaera feae Attems, 1935: 128 (first description)

Indosphaera feae — Jeekel 2001: 15 (list)

— Indosphaera curiosa Attems, 1936: 183 (first descrip.)

Indosphaera curiosa :— Rajulu 1970: 128 (list), syn-

onymized under I feae by Jeekel 2001: 15.

Remark: I. curiosa is an objective synonym of I feae, a

name not used in the second proposal of Indosphaera as

a new genus in 1936.

Types: M and F syntypes.

Habitat: India: Nemotha and Silcuri, Cachar, Assam.

Deposited: NHMW 2225 (Stagl & Schiller, in prep.).

Indosphaera fortis Attems, 1936

Indosphaera fortis Attems, 1936: 186 (first description)

Indosphaera fortis — Jeekel 2001: 15 (list).

Types: F holotype (dry).

Habitat: Burma: Southern Shan States.

Deposited: NHMW 2224 (Stagl & Schiller, in prep.).

Kopho spinier a Attem s. 1935

Attems, 1935: 128 (first description); Attems 1936: 153

(description); Attems 1943: 62 (list); Jeekel 1971: 26 (list);

Jeekel 1974: 46 (classification); Hoffman 1976: 119 (list);

Hoffman 1980: 63 (list); Jeekel 2001: 15 (list); Mauries

2001: 141 (list); Wesener & VandenSpiegel 2009: 551

(list); Wesener 2015 (key, redescription).

Description: Anterior telopod with four podomeres dis¬

tal to syncoxite. Telopoditomere 2 with a large, curved

process forming a clamp with simple telopoditomeres 3

and 4. Posterior telopod with four telopoditomeres,

telopoditomere 4 quite large. Posterior telopod lacking any

membranous processes or sclerotized teeth. Telopodito¬

meres 3 and 4 each with a membranous ledge, spines on¬

ly on telopoditomere 4; general shape similar to Zephro-

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Species Catalogue Sphaerotheriida

nia. Antennomere 6 strongly flattened. Legs slender, with

a single apical spine. Anal shield with single black lock¬

ing carina, in some species reduced. 3CM5 mm long. Ter-

gites glabrous. Might fall into synonymy under Zephro¬

nia.

Type-species: Kophosphaera politissima Attems, 1935.

Species included (7)

Kophosphaera brevilamina Attems, 1936 (NE India)

Kophosphaera devolvens Attems, 1936 (NE India)

Kophosphaera excavata (Butler, 1874) (NE India, Nepal)

Kophosphaera mammifera Attems, 1936 (NE India)

Kophosphaera martensi Wesener, 2015 (Nepal)

Kophosphaera politissima Attems, 1935 (NE India,

Nepal)

Kophosphaera shivapuri Wesener, 2015 (Nepal)

Kophosphaera brevilamina Attems, 1936

Kophosphaera brevilamina Attems, 1936: 191 (first de¬

scription)

Kophosphaera brevilamina :— Rajulu 1970: 129 (list)

Kophosphaera brevilamina :— Jeekel 2001: 15 (list)

Kophosphaera brevilamina.— Sanyal et al. 2012: 865 (list)

Kophosphaera brevilamina — Wesener 2015: 309 (key).

Types: 3 M? syntypes.

Habitat: India: Siliguri, North Bengal; Korseong, Eastern

Himalayas.

Deposited: NHMW 100/191 (Stagl & Schiller, in prep.).

Kophosphaera devolvens Attems, 1936

Kophosphaera devolvens Attems, 1936: 190 (first descrip.)

Kophosphaera devolvens :— Rajulu 1970: 129 (list)

Kophosphaera devolvens :— Jeekel 2001: 15 (list)

Kophosphaera devolvens :— Sanyal et al. 2012: 865 (list)

Kophosphaera devolvens :— Wesener 2015: 309 (key).

Types: 3 M? syntypes.

Habitat: India: Gangtok, Sikkim; Soreil, Darjeeling dis¬

trict; Darjeeling, Eastern Himalayas.

Deposited: NHMW 2222, 2223 (Stagl & Schiller, in

prep.).

Kophosphaera excavata (Butler, 1874)

Zephronia excavata Butler, 1874: 185 (first description)

Zephronia excavata :— Preudhomme de Borre 1884: 24

(list)

Zephronia excavata — Pocock 1899: 466 (description)

Zephronia excavata — Attems 1914: 145 (list)

Kophosphaera excavata — Attems 1936: 187 (new comb.)

Kophosphaera excavata :— Jeekel 2001: 15 (list)

Kophosphaera excavata :— Wesener 2015: 309 (key, re¬

description).

Remarks: Attems redescription probably not identical with

original (Jeekel 2001: 15).

Types: Lectotype, 3 syntypes (apparently lost).

Habitat: India: Sikkim (Type); Dikrang Valley, Darrang

distr., Shillong, Khasi Hills, Assam; Nepal: Chitlong.

Deposited: NHML BM1874.51.

Kophosphaera mammifera Attems, 1936

Kophosphaera excavata, ssp. mammifera Attems, 1936:

188 (first description)

Kophosphaera excavata, ssp. mammifera :— Jeekel 2001:

16 (list)

Kophosphaera excavate (sic!), mammifera — Sanyal et al.

2012: 865 (list)

Kophosphaera mammifera:— Wesener 2015: 311 (new sta¬

tus).

Types: 10 M and F syntypes.

Habitat: India: Sureil, Darjeeling distr.; Eastern Hi¬

malayas; Khasi Hills, Assam.

Deposited: NHMW 2221 (Stagl & Schiller, in prep.).

Kophosphaera martensi Wesener, 2015

Kophosphaera martensi Wesener, 2015: 316 (first descrip¬

tion).

Types: Holotype

Habitat: Nepal: Kaski District, Dhumpus.

Deposited: SMF.

Kophosphaera politissima Attems, 1935

Kophosphaera politissima Attems, 1935: 128 (first de¬

scription)

Kophosphaerapolitissima:— Attems 1936: 180 (descrip.)

Kophosphaera politissima :— Rajulu 1970: 129 (list)

Kophosphaera politissima :— Jeekel 2001: 16 (list)

Kophosphaera politissima:— Sanyal et al. 2012: 865 (list)

Kophosphaera politissima :— Wesener 2015: 309 (re¬

description).

Types: 4 M and F syntypes.

Habitat: India: Dobie Jhora, Kurseong; Sureil, Darjeeling

distr., Eastern Himalayas; Nepal.

Deposited: NHMW 2220 (Stagl & Schiller, in prep.).

Kophosphaera shivapuri Wesener, 2015

Kophosphaera shivapuri Wesener, 2015: 313 (first descr.).

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Types: Holotype

Habitat: Nepal: Kathmandu District, Shivapuri Mountain.

Deposited: SMF.

Leptotelopus Silvestri. 1897

Leptotelopus Silvestri, 1897c: 645 (first description);

Jeekel 1971: 26 (list); Jeekel 1974: 46 (classification);

Hoffman 1980: 63 (list); Jeekel 2001: 16 (list); Mauries

2001: 141 (list); Wesener & VandenSpiegel 2009: 551

(list).

Synonyms:

— Lophozephronia Attems, 1936: 152, synonymized by

Jeekel 1971: 26. Attems, 1943: 61; Attems, 1953: 155;

Jeekel, 1971: 26.

Type species'. Zephronia crepitans Pocock, 1890.

Description: Male legs in posterior half uniquely modi¬

fied, with enlarged coxa and prefemur, and greatly elon¬

gated, but very slender femur, postfemur, tibia and tarsus.

Tarsus with fringes of hair at lateral and mesal margins.

Anterior telopods with three podomeres distal to syncox-

ite. Telopoditomere 3 small, forming a clamp with a

process of telopoditomere 2. Posterior telopods with four

telopoditomeres. A large and ridged plate located on

telopoditomere 4 might represent a unique stridulation or¬

gan. 15 mm long.

Type-species: Zephronia crepitans Pocock, 1890.

Monotypic

Leptotelopus crepitans (Pocock, 1890)

Zephronia crepitans Pocock, 1890b: 392 (first description)

Leptotelopus crepitans.— Silvestri, 1897c: 645 (type of

new genus)

Zephronia crepitans :— Pocock 1899: 471 (list)

Zephronia crepitans :— Attems 1914: 145 (list)

Lophozephronia crepitans.— Attems, 1936: 183 (new com¬

bination)

Lophozephronia crepitans :— Jeekel 2001: 16 (list)

Leptotelopus crepitans :— Jeekel 2001: 16 (list)

Leptotelopus crepitans — Mauries 2001: 141 (list).

Types: 5 M and F syntypes.

Habitat: Burma: Rangoon.

Deposited: NHML BM1889.12.20.13-16.

Zephronia Grav. 1832

(Figs 11D-H)

Zephronia Gray, 1832: 796; Gervais 1837: 42; Newport

1844: 265; Gervais 1844: 61; Gervais 1847: 75; Butler

1873: 178; Pocock 1894: 326; Pocock 1895: 413; Silvestri

1896b: 205; Pocock 1899: 270; Flower 1901: 7; Attems

1914: 142; Attems 1926: 120; Attems 1935: 129; Attems

1936: 153; Attems 1943: 63; Attems 1953: 155; Jeekel

1971: 29 (list); Jeekel 1974: 46 (classification); Hoffman

1976: 120 (comments); Hoffman 1980: 64 (list); Jeekel

2001: 17 (list); Wesener & VandenSpiegel 2009: 551 (mor¬

phological phylogenetic analysis); Wesener et al. 2010:

1188 (molecular phylogenetic analysis); Golovatch et al.

2012: 283 (redescription); Peeters & De Greef 2015: 3

(predation by ants).

Keys: Pocock 1890; Pocock 1899; Attems 1936.

Description: Anterior telopods with four podomeres dis¬

tal to syncoxite. Telopoditomere 2 with a large, curved

process forming a clamp with simple telopoditomeres 3

and 4. Posterior telopod with four telopoditomeres. Im¬

movable finger slender, curved apically, towards telopodit¬

omeres 3 and 4 always with two membranous ledges.

Telopoditomere 3 also with a membranous ledge and a row

of large, crenulated, sclerotized teeth, much larger than

telopoditomere 4. Telopoditomere 4 with sclerotized ledge

and few sclerotized speines. Antennae flattened laterally,

axe-shaped. 18-50 mm long. Tarsi often with several api¬

cal spines.

Type-species. Zephronia ovalis Gray, 1832.

Species and sub-species included (39)

Zephronia alticola alticola Attems, 1936 (NE India)

Zephronia alticola bengalica Attems, 1936 (NE India)

Zephronia anthracina Pocock, 1895 (Malaysia, Perak)

Zephronia butleri Olliff, 1882 (North Borneo)

Zephronia clivicola Pocock, 1890 (Burma)

Zephronia comotti Pocock, 1890 (Burma)

Zephronia dawydoffi Attems, 1953 (Cambodia)

Zephronia debilis Attems, 1936 (NE India)

Zephronia densipora Attems, 1936 (NE India)

Zephronia disparipora Attems, 1936 (NE India)

Zephronia dollfusi Pocock, 1895 (Vietnam)

Zephronia doriae Pocock, 1890 (Burma)

Zephronia feae Pocock, 1890 (Burma)

Zephronia floweri Hirst, 1907 (Singapore)

Zephronia formosa Pocock, 1890 (Burma)

Zephronia gestri Pocock, 1890 (Burma)

Zephronia hirsti Jeekel 2001 (Malaysia)

Zephronia hirta Attems, 1936 (NE India)

Zephronia hysophila Attems, 1936 (NE India)

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Species Catalogue Sphaerotheriida

Zephronia impunctata Pocock, 1895 (Malaysia)

Zephronia inferior Attems, 1936 (Burma)

Zephronia juvenis Attems, 1936 (NE India)

Zephronia lignivora Attems, 1936 (NE India)

Zephronia maculatum Butler, 1874 (NE India)

Zephronia manca Attems, 1936 (Vietnam, NE India)

Zephronia montana (Karsch, 1881) (‘Himalaya’)

Zephronia nepalensis Wesener, 2015 (Nepal)

Zephronia nigrinota Butler, 1872 (NE India)

Zephronia ovalis Gray, 1832 (Vietnam)

Zephroniapellita Attems, 1935 (Sumatra)

Zephronia profuga Attems, 1936 (China, Hongkong)

Zephronia rid/eyi Hirst, 1907 (Malaysia)

Zephronia semilaevis Pocock, 1890 (Bunna)

Zephronia siamensis Hirst, 1907 (Thailand)

Zephronia specularis Attems, 1936 (NE India)

Zephronia tigrina Butler, 1872 (NE India)

Zephronia tigrinoides Attems, 1936 (NE India)

Zephronia tumida Butler, 1882 (NE India, Burma)

Zephronia viridescens Attems, 1936 (Burma)

Remark: The name Zephronia has been used to cover

many generic concepts; its concept was not fixed until the

redescription of the type species (Golovatch et al. 2012).

However, a revision of numerous described species is nec¬

essary to evaluate whether or not they fall into the rede¬

fined genus.

Zephronia alticola alticola Attems, 1936

Zephronia alticola Attems, 1936: 176 (first description)

Zephronia alticola -- Rajulu 1970: 128 (list)

Zephronia alticola.— Jeekel 2001: 17 (list)

Zephronia alticola.— Sanyal et al. 2012: 865 (list).

Types: M and F syntypes.

Habitat: India: Sureil, Darjeeling distr., Eastern Himalayas;

Rotung, Assam.

Deposited: NHMW 2235 (Stagl & Schiller, in prep.).

Zephronia alticola bengalica Attems, 1936

Zephronia alticola, ssp. bengalica Attems, 1936: 177 (first

description)

Zephronia alticola, ssp. bengalica :— Jeekel 2001: 17 (list).

Types: M holotype.

Habitat: India: Buxa Road, North Bengal.

Deposited: NHMW 2236.

Zephronia anthracina Pocock, 1895

Zephronia anthracina Pocock, 1895: 413 (first descrip.)

Zephronia anthracina :— Hirst, 1907: 215 (list)

Zephronia anthracina:— Attems, 1914: 144 (list)

Zephronia anthracina :— Wang 1967: 384 (list)

Zephronia anthracina :— Jeekel 2001: 18 (list).

Types: Holotype? of unknown sex.

Habitat: Malaysia: Perak.

Deposited: NHML (Online Database).

Zephronia butleri Olliff, 1882

Zephronia Butleri Olliff, 1882: 29 (first description)

Zephronia Butleri :— Preudhomme de Borre 1884: 23 (list)

Zephronia butleri :— Jeekel 2001: 18 (list).

Types: M and F syntypes.

Habitat: North Borneo: Sandakan.

Deposited: NHML (not listed as type, Online Database).

Zephronia clivicola Pocock, 1890

Zephronia clivicola Pocock, 1890b: 386 (first description)

Zephronia clivicola:— Pocock 1899: 469 (list)

Zephronia clivicola:— Attems 1914: 145 (list)

Zephronia clivicola :— Attems, 1936: 170 (new locality)

Zephronia clivicola :— Weidner 1960: 78 (list)

Zephronia clivicola:— Jeekel 2001: 18 (list).

Types: M and F syntypes.

Habitat: Burma: Carin Cheba or Biapo (villages of Puepoli

and MeLeleo); Second Defile, River Irrawaddy.

Deposited: NHML (Online Database), 2 paratypes ZMUH

(Weidner 1960: 78).

Zephronia comotti Pocock, 1890

Zephronia comotti Pocock, 1890b: 391 (first description)

Zephronia comotti — Pocock 1899: 471 (list)

Zephronia comotti :— Attems 1914: 145 (list)

Zephronia comotti :— Attems 1936: 182 (list)

Zephronia comotti — Jeekel 2001: 18 (list).

Types: M holotype?

Habitat: Burma: Minhla.

Deposited: NHML?

Zephronia dawydoffi Attems, 1953

(Fig. 11D)

Zephronia dawydoffi Attems, 1953: 156 (first description)

Zephroniacambodjana [sic!]:—Attems 1953: pi. 3 (error)

Zephronia dawydoffi :— Jeekel 2001: 18 (list)

Zephronia dawydoffi- Likhitrakam et al. 2015: 177 (list).

Remark: In the captions of the illustrations the name of

the new species is Zephronia cambodjana , an obvious

clerical error (Jeekel 2001).

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Types: Syntypes.

Habitat: Cambodia: Ream, Sre Umbell; Kampot, Kam-

pong Speu.

Deposited: NHMW 2241 (Stagl & Schiller, in prep.).

Zephronia debilis Attems, 1936

Zephronia debilis Attems, 1936: 172 (first description)

Zephronia debilis -- Rajulu 1970: 127 (list)

Zephronia debilis -- Jeekel 2001: 18 (list)

Zephronia debilis -- Sanyal et al. 2012: 865 (list).

Types: F holotype.

Habitat: India: Darjeeling; Eastern Himalayas.

Deposited: NHMW 3956 (Stagl & Schiller, in prep.).

Zephronia densipora Attems, 1936

Zephronia densipora Atterns, 1936: 171 (first description)

Zephronia densipora:— Rajulu 1970: 127 (list)

Zephronia densipora:— Jeekel 2001: 18 (list).

Types: M holotype.

Habitat: India: Elephant Falls, Shillonp; Khasi Hills, As¬

sam.

Deposited: NHMW 2237 (Stagl & Schiller, in prep.).

Zephronia disparipora Attems, 1936

Zephronia disparipora Attems, 1936: 171 (first descript.)

Zephronia disparipora:— Rajulu 1970: 127 (list)

Zephronia disparipora:— Jeekel 2001: 18 (list).

Types: M holotype without telopods.

Habitat: India: Kobo, Assam.

Deposited: Indian Museum Calcutta? (now Zoological

Survey of India, Kolkata).

Zephronia dollfusi Pocock, 1895

Zephronia dollfusi Pocock, 1895: 413 (first description)

Zephronia dollfusi :— Attems 1914: 145 (list)

Zephronia dollfusi :— Attems 1936: 182 (list)

‘ Zephronia ’ dollfusi :— Golovatch 1983 (list)

Zephronia dollfusi — Jeekel 2001: 18 (list)

Zephronia dollfusi :— Enghoff et al. 2004: 32 (list).

Types: Holotype of unknown sex.

Habitat: Vietnam: Cochin China.

Deposited: NHML.

Zephronia doriae Pocock, 1890

Zephronia doriae Pocock, 1890a: 79 (first description)

Zephronia doriae — Pocock 1890b: 385 (list)

Zephronia doriae :— Pocock 1899: 469 (list)

Zephronia doriae :— Attems 1914: 145 (list)

Zephronia doriae — Attems 1936: 182 (list.

Zephronia doriae :— Jeekel 2001: 18 (list).

Types: M and F syntypes.

Habitat: Burma: Kachin Hills, East of Bhamo.

Deposited: NHML (Online Catalogue).

Zephronia feae Pocock, 1890

Zephronia feae Pocock, 1890a: 80 (first description)

Zephronia feae — Pocock 1890b: 385 (new locality)

Zephronia feae — Pocock 1899: 469 (list)

Zephronia feae — Attems 1914: 145 (list)

Zephronia feae — Attems 1936: 182 (list)

Zephronia feae — Weidner 1960: 78 (list)

Zephronia feae — Jeekel 2001: 18 (list).

Types: M and F syntypes.

Habitat: Burma: Teinzo, Moolay River (Type); Kachin

Cauri, East of Bhamo.

Deposited: NHML (Online Catalogue), 5 paratypes

ZMUH (Weidner 1960: 78).

Zephronia flowed Hirst, 1907

Zephronia flowed Hirst, 1907: 217 (first description)

Zephronia flowed.— Attems 1914: 145 (list)

Zephronia flowed :— Jeekel 2001: 19 (list)

Zephronia flowed — Decker 2013: 3 (list).

Type: Two F syntypes.

Habitat: Singapore.

Deposited: NHML (Online Cat., author listed as Pocock).

Zephronia formosa Pocock, 1890

Zephronia formosa Pocock, 1890b: 387 (first description)

Zephronia formosa:— Pocock 1899: 470 (description)

Zephronia formosa — Attems 1914: 145 (list)

Zephronia formosa:— Attems 1936: 182 (list)

Zephronia formosa:— Weidner 1960: 79 (list)

Zephronia formosa:— Jeekel 2001: 19 (list).

Types: M and F syntypes.

Habitat: Burma: Carin Ghecu (village of Thao); Carin

Cheba or Biapo (villages of Puepoli and Meteleo).

Deposited: NHML (Online Catalogue, unclear if types),

2 paratypes, ZMUH (Weidner 1960: 79).

Zephronia gestri Pocock, 1890

Zephronia gestri Pocock, 1890b: 390 (first description)

Zephronia gestri — Pocock 1899: 470 (list)

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Species Catalogue Sphaerotheriida

Zephronia gestri — Attems 1914: 145 (list)

Zephronia gestri. — Attems 1936: 182 (list)

Zephronia gestri Jeekel 2001: 19 (list).

Types: M holotype.

Habitat: Burma: Thagata on the slope of Mt. Mooleyit.

Deposited: NHML?

Zephronia hirsti Jeekel, 2001

Zephronia hirsti Jeekel 2001:19 (first description)

— Zephronia rugulosa Hirst, 1907: 215 (first description)

Zephronia rugulosa.- Attems, 1914: 146 (list) NOT

Zephronia rugulosa Gervais, 1847: 62

(= Sphaerotherium rugulosum Brandt, 1841: 363), and

NOT Zephronia rugulosa Butler, 1872 = Arthrosphaera

ruginosa Jeekel 2001 (synonymized Jeekel 2001: 19).

Types: 3 M syntypes.

Habitat: Malaysia: Pahang; Kelantan.

Deposited: NHML (Online Catalogue, unclear if types).

Zephronia hirta Attems, 1936

Zephronia hirta Attems, 1936: 179 (first description)

Zephronia hirta .— Rajulu 1970: 128 (list)

Zephronia hirta :— Jeekel 2001: 19 (list)

Zephronia hirta .— Sanyal et al. 2012: 865 (list).

Types: F holotype.

Habitat: India: Sureil, Darjeeling distr., Eastern Himalayas.

Deposited: Indian Museum Calcutta? (now Zoological

Survey of India, Kolkata).

Zephronia hysophila Attems, 1936

Zephronia hysophila Attems, 1936: (first description)

Zephronia hysophila:— Rajulu 1970: 128 (list)

Zephronia hysophila :— Jeekel 2001: 19 (list).

Types: F holotype (missing leg 2).

Habitat: India: Dikrang Valley, Darrang distr. Assam.

Deposited: Indian Museum Calcutta? (now Zoological

Survey of India, Kolkata).

Zephronia impunctata Pocock, 1895

Zephronia impunctata Pocock, 1895: 413 (first descrip.)

Zephronia impunctata :— Flower 1901: 8 (list)

Zephronia impunctata :— Hirst 1907: 215 (description)

Zephronia impunctata — Attems 1914: 146 (list)

Zephronia impunctata :— Attems 1936: (new locality)

Zephronia impunctata :— Wang 1967: 384 (list)

Zephronia impunctata :— Jeekel 2001: 19 (list).

Types: holotype of unknown sex.

Habitat: Malaysia: Penang.

Deposited: NHML (Online Catalogue).

Zephronia inferior Attems, 1936

Zephronia inferior Attems, 1936: 178 (first description)

Zephronia inferior— Jeekel 2001: 19 (list).

Types: M and F syntypes.

Habitat: Burma: Moulmein.

Deposited: NHMW, unclear if types (Stagl & Schiller, in

prep.).

Zephronia juvenis Attems, 1936

Zephronia juvenis Attems, 1936: 171 (first description)

Zephronia juvenis — Rajulu 1970: 127 (list)

Zephronia juvenis :— Jeekel 2001: 19 (list).

Types: F holotype.

Habitat: India: Darrang distr., Assam.

Deposited: NHMW, unclear if types, (Stagl & Schiller, in

prep.).

Zephronia lignivora Attems, 1936

Zephronia lignivora Attems, 1936: 174 (first description)

Zephronia lignivora :— Rajulu 1970: 127 (list)

Zephronia lignivora :— Jeekel 2001: 19 (list).

Types: M and F syntypes.

Habitat: India: Kobo; Geku, Assam.

Deposited: NHMW 2238 (Stagl & Schiller, in prep.).

Zephronia maculata (Butler, 1874)

Sphaerotherium maculatum Butler, 1874: 186 (first de¬

scription)

Sphaerotherium maculatum — Preudhomme de Borre

1884: 27 (list)

Zephronia maculata :— Pocock 1899: 467 (description)

Sphaerotherium maculatum:— Attems 1914: 141 (list)

Sphaerotherium maculatum — Attems 1936: 192 (list)

Zephronia maculata :— Jeekel 2001: 20 (list).

Types: F holotype.

Habitat: India: Sikkim.

Deposited: NHML (Online Catalogue).

Zephronia manca Attems, 1936

Zephronia manca Attems, 1936: 169 (first description)

Zephronia manca:— Rajulu 1970: 127 (list)

‘ Zephronia ’ manca:— Golovatch 1983 (list)

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Zephronia manca — Enghoff et al. 2004: 32 (list).

Zephronia manca.— Jeekel 2001: 20 (list)

Zephronia manca — Sanyal et al. 2012: 865 (list).

Types: M and F syntypes.

Habitat: Vietnam: Dran; Dalat, FangbianProv., South An-

nam. India: Dhobie Jhora,

Kurseong, Eastern Himalayas.

Deposited: NHMW 2239 (Stagl & Schiller, in prep.).

Zephronia montana (Karsch, 1881)

Sphaeropoeus montanus Karsch, 1881: 31 (first descrip¬

tion)

Zephronia montana — Preudhomme de Borre: 25 (list)

Sphaeropoeus montanus :— Attems 1914: 143 (list)

Sphaeropoeus montanus — Attems 1936: 192 (list)

Sphaeropoeus montanus — Moritz & Fischer 1974: 369

(list)

Sphaeropoeus montanus — Jeekel 2001: 24 (list)

Zephronia montana :— Wesener 2015: 302 (new combina¬

tion).

Types: F holotype.

Habitat: “Himalaya”

Deposited: ZMB 55.

Zephronia nepalensis Wesener, 2015

Zephronia nepalensis Wesener, 2015: 305 (first descrip.).

Types: M holotype, 1 M, 1 F paratypes.

Habitat: Nepal, Ilam District, Mai Pokhari.

Deposited: SMF (holotype), ZFMK MYR4173 & 4172

(paratypes).

Zephronia nigrinota Butler, 1872

Zephronia nigrinota Butler, 1872: 356 (first description)

Zephronia nigrinota :— Preudhomme de Borre 1884: 25

(list)

Zephronia nigrinota :— Pocock 1899: 468 (description,

new locality)

Zephronia nigrinota :— Attems 1914: 146 (list)

Zephronia nigrinota — Attems 1936: 173 (description)

Zephronia nigrinota :— Rajulu 1970: 127 (list)

Zephronia nigrinota :— Jeekel 2001: 20 (list)

“ Zephronia ” nigrinota :— Beron 2008: 169 (list)

Zephronia nigrinota [sic!]:— Sanyal et al. 2012: 865 (list)

— Zephronia laevissima Butler, 1874: 185 (first descrip.),

synonymized with Z nigrinota by Pocock 1899: 356

Zephronia laevissima — Preudhomme de Borre 1884: 24

(list)

Zephronia laevissima :— Pocock 1899: 356 (synonymized

with Z nigrinota)

Zephronia laevissima — Attems 1914: 146 (list)

Zephronia laevissima :— Attems 1936: 182 (list)

— Sphaerotherium politum Butler, 1874: 186 (first de¬

scription). synonymized with Z nigrinota by Pocock

1899:356

Sphaerotherium politum :— Preudhomme de Borre 1884:

27 (list)

Sphaerotherium politum — Pocock 1899: 356 (syn¬

onymized with Z. nigrinota)

Sphaerotherium politum — Attems 1914: 141 (list)

Sphaerotherium politum — Attems 1936: 193 (list).

Types: M and F syntypes.

Habitat: India: Sikkim Assam): Aideo, Assam; Darjeeling,

Jorpokrei, Eastern

Himalayas.

Deposited: NHMF (Online Catalogue).

Zephronia ovalis Gray, 1832

Zephronia ovalis Gray, 1832: 796 (first description)

Zephronia ovalis :— Gervais 1837: 42 (list)

Zephronia ovalis :— Newport 1844: 264 (new locality)

Sphaerotherium ovale - CF Koch 1847: 99 (list)

Zephronia ovalis :— Butler 1873: 180 (list)

Zephronia ovalis :— Olliff 1882: 30 (description)

Zephronia ovalis :— Preudhomme de Borre 1884: 25 (list)

Zephronia ovalis :— Attems 1914: 146 (list)

Zephronia ovalis :— Jeekel 2001: 20 (list)

Zephronia ovalis — Golovatch et al. 2012: 286 (redescrip.)

Zephronia ovalis — Wesener 2014a: 204 (morphological

phylogenetic analysis).

Types: M neotype (designated in Golovatch et al. 2012:

286).

Habitat: Vietnam, Cat Tien National Park.

Deposited: ZMUM.

Zephronia pellita Attems, 1935

Zephronia pellita Attems, 1935: 141 (first description)

Zephronia pellita - Jeekel 2001: 21 (list).

Remarks: The distribution of this species lies outside the

area of distribution of the other ZQphronia species.

Types: M holotype.

Habitat: Indonesia: Sumatra, Indragiri, Sungei Falah.

Deposited: NHMW (non-type material?) (Stagl & Schiller,

in prep.).

Zephronia profuga Attems, 1936

Zephronia profuga Attems, 1936: 181 (first description)

Zephronia profuga :— Jeekel 2001: 21 (list).

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Species Catalogue Sphaerotheriida

Types: F holotype.

Habitat: China: Hongkong.

Deposited: NHMW 2240 (Stagl & Schiller, in prep.).

Zephronia ridleyi Hirst, 1907

Zephronia ridleyi Hirst, 1907: 216 (first description)

Zephronia ridleyi -- Attems 1914: 146 (list)

Zephronia ridleyi.— Jeekel 2001: 21 (list).

Types: M holotype.

Habitat: Malaysia: Selangor.

Deposited: NHML (Online Catalogue).

Zephronia semilaevis Pocock, 1890

Zephronia semilaevis Pocock, 1890b: 388 (first descrip.)

Zephronia semilaevis :— Pocock 1899: 470 (description)

Zephronia semilaevis .— Hirst 1907: 215 (list)

Zephronia semilaevis — Attems 1914: 146 (list)

Zephronia semilaevis — Attems 1936: 182 (list)

Zephronia semilaevis :— Jeekel 2001: 21 (list).

Types: M and F syntypes.

Habitat: Burma: S. Tenasserim; Malewoon (Tenasserim).

Deposited: NHML (Online Catalogue); 5 paratypes,

ZMUH (Weidner 1960: 79).

Zephronia siamensis Hirst, 1907

Zephronia siamensis Hirst, 1907: 218 (first description)

Zephronia siamensis.— Attems 1914: 147 (list)

Zephronia siamensis Attems 1936: 182 (list)

Zephronia siamensis Jeekel 2001: 21 (list)

Zephronia siamensis Golovatch et al. 2012: 276 (genet¬

ic barcode).

Types: M and F syntypes.

Habitat: Thailand: Kosichang; Chantaboon.

Deposited: NHML (Online Catalogue).

Zephronia specularis Attems, 1936

Zephronia specular is Attems, 1936: 174 (first description)

Zephronia specularis :— Rajulu 1970: 127 (list)

Zephronia specularis :— Jeekel 2001: 21 (list).

Types: M and F syntypes.

Habitat: India: Dafla Hills, Assam.

Deposited: NHMW 2233 (Stagl & Schiller, in prep.).

Zephronia tigrina Butler, 1872

Zephronia tigrina Butler, 1872: 356 (first description)

Zephronia tigrina :— Preudhomme de Borre 1884: 25 (list)

Zephronia tigrina :— Pocock 1899: 466 (description)

Zephronia tigrina :— Attems 1914: 147 (list)

Zephronia tigrina — Jeekel 2001: 21 (list).

Types: M and F syntypes.

Habitat: “East Indies”. India: Darjeeling.

Deposited: NHML (Online Catalogue).

Zephronia tigrinoides Attems, 1936

Zephronia tigrinoides Attems, 1936: 177 (first description)

Zephronia tigrinoides :— Rajulu 1970: 128 (list)

Zephronia tigrinoides :— Jeekel 2001: 21 (list)

Zephronia tigrinoides — Sanyal et al. 2012: 865 (list).

Types: M holotype.

Habitat: India: Sukna, Darjeeling distr., Eastern Hi¬

malayas.

Deposited: Indian Museum Calcutta? (now Zoological

Survey of India, Kolkata).

Zephronia tumida Butler, 1882

Zephronia tumida Butler, 1882: 196 (first description)

Zephronia tumida :— Preudhomme de Borre 1884: 25 (list)

Zephronia tumida :— Pocock 1899: 465 (description)

Zephronia tumida — Attems 1914: 147 (list)

Zephronia tumida — Attems 1936: 175 (description)

Zephronia tumida — Rajulu 1970: 128 (list)

Zephronia tumida :— Jeekel 2001: 21 (list)

Zephronia tumida var. nigrinota [sic!]:— Sanyal et al.

2012: 865 (list)

Zephronia tumida :— Wesener 2015: 303 (redescription).

Types: M holotype.

Habitat: India: N. Assam (Type); India: Harmutti, base of

Dafla Hills; Sibsagar; Dikrang Valley, Darrang distr. As¬

sam; Darjeeling, Eastern Himalayas. Myanmar: Bunna:

Tavoy, Moti Ram, lower Burma, Kamaing, Myitkyina dis¬

tr., upper Burma.

Deposited: NHML BM1882.11.

Zephronia viridescens Attems, 1936

Zephronia viridescens Attems, 1936: 180 (first descrip.)

Zephronia viridescens :— Jeekel 2001: 22 (list)

Zephronia cf. viridescens :— Wongthamwanich et al.

2012b: 913 (ecology)

Zephronia cf. viridescens :— Sukteeka & Thanee 2015: 18

(ecology).

Types: 1 M and 1 F syntypes.

Habitat: Burma: Tavoy.

Deposited: Indian Museum Calcutta (now Zoological Sur¬

vey of India, Kolkata).

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Tribe Sphaeropoeini Brolemann, 1913

Name adapted from Sphaeropoeidae Brolemann, 1913: 79;

Hoffman 1976: 119; Hoffman 1980: 63; Jeekel 2001: 22.

— Priobelini Hoffman, 1976, synonymized by Mauries

2001: 140.

Sphaerobelum Verhoeff. 1924

Sphaerobelum Verhoeff, 1924: 59 (first description); At-

tems 1926: 120 (list); Attems 1928: 203 (list); Attems

1936: 153 (list); Attems, 1938: 200 (new species); Attems

1943: 62 (list); Attems 1953: 155 (new species); Jeekel

1971: 28 (list); Jeekel 1974: 46 (classification); Hoffman

1976: 119 (list); Hoffman 1980: 63 (list); Jeekel 2001: 17

(list); Mauries 2001: 141 (list); Wesener & VandenSpiegel

2009: 551 (list); Wongthamwanich et al. 2012a: 34 (new

species).

Description: Anterior telopod with 3 or 4 podomeres dis¬

tal to syncoxite. Second telopoditomere with a curved

process, more distally located telopoditomeres usually

simple. Posterior telopods with four telopoditomeres, im¬

movable finger apically massively enlarged, a unique char¬

acter. 20^15 mm long. Tarsi with single apical spine. Sur¬

face variable, hairy to shiny and glabrous.

Type-species: Sphaerobelum clavigerum Verhoeff, 1924.

Species included (5)

Sphaerobelum bicorne Attems, 1938 (Vietnam, Bana)

Sphaerobelum clavigerum Verhoeff, 1924 (Vietnam,

Tonkin)

Sphaerobelum hirsutum Verhoeff, 1924 (Vietnam, Tonkin)

Sphaerobelum separatum Attems, 1953 (Vietnam, Tonkin,

Fan-Si-Pan)

Sphaerobelum truncatum Wongthamwanich, 2012 (Thai¬

land, Nan)

Sphaerobelum bicorne Attems, 1938

Sphaerobelum bicorne Attems, 1938: 200 (first descrip.)

Sphaerobelum bicorne.— Enghoff et al. 2004: 32 (list)

Sphaerobelum bicorne :— Jeekel 2001: 17 (list)

Sphaerobelum bicorne :— Wongthamwanich et al. 2012a:

34 (key).

Types: M and F syntypes.

Habitat: Vietnam: Bana (Tourane, Annam).

Deposited: NHMW 2196 (Stagl & Schiller, in prep.).

Sphaerobelum clavigerum Verhoeff, 1924

Sphaerobelum clavigerum Verhoeff, 1924: 65 (first de¬

scription)

Sphaerobelum clavigerum :— Attems 1936: 192 (list)

Sphaerobelum clavigerum :— Moritz & Fischer 1973: 368

(list)

Sphaerobelum clavigerum :— Golovatch 1983 (list)

Sphaerobelum clavigerum :— Jeekel 2001: 17 (list)

Sphaerobelum clavigerum:— Enghoff et al. 2004: 32 (list)

Sphaerobelum clavigerum — Wongthamwanich et al.

2012a: 34 (key)

Sphaerobelum clavigerum :— Wesener 2014a: 204 (mor¬

phological phylogenetic analysis).

Types: Holotype M, syntype.

Habitat: Vietnam: Tonkin.

Deposited: Holotype ZMB 5747 (Moritz & Fischer 1973:

368); syntype HNHM (Korsos 1983: 117)

Sphaerobelum hirsutum Verhoeff, 1924

Sphaerobelum hirsutum Verhoeff, 1924: 66 (first descrip¬

tion)

Sphaerobelum hirsutum :— Attems, 1936: 192 (list)

Sphaerobelum hirsutum — Golovatch 1983 (list)

Sphaerobelum hirsutum — Jeekel 2001: 17 (list)

Sphaerobelum hirsutum — Enghoff et al. 2004: 32 (list)

Sphaerobelum hirsutum :— Wongthamwanich et al. 2012a:

34 (key).

Types: 2 M, 2 F syntypes.

Habitat: Vietnam: Tonkin.

Deposited: Unknown, at least telopods at ZSM.

Sphaerobelum separatum Attems. 1953

Sphaerobelum separatum Attems, 1953: 155 (first descrip¬

tion)

Sphaerobelum separatum — Golovatch 1983 (list)

Sphaerobelum separatum :— Jeekel 2001: 17 (list)

Sphaerobelum separatum :— Enghoff et al. 2004: 32 (list)

Sphaerobelum separatum — Wongthamwanich et al.

2012a: 34 (key).

Types: M holotype.

Habitat: Vietnam: Fan-Si-Pan, Tonkin.

Deposited: NHMW 2227 (Stagl & Schiller, in prep.).

Sphaerobelum truncatum Wongthamwanich, 2012

Sphaerobelum truncatum Wongthamwanich et al. 2012a:

34 (first description)

Sphaerobelum truncatum — Wesener 2014a: 204 (morpho¬

logical phylogenetic analysis).

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Species Catalogue Sphaerotheriida

Types: M holotype, 6M,6F paratypes.

Habitat: Thailand, Nan Province, Na Rai Luang Sub-dis¬

trict, Pang Hi Village.

Deposited: CUMZ (holotype, 8 paratypes); FMNH (2

paratypes); ZFMK (2 paratypes).

Prionobelum Verhoeff. 1924

Prionobelum VerhoefF, 1924: 58 (first description); Attems

1926: 120 (list); Attems 1935: 131 (new species); Attems

1936: 153 (list); Attems 1943: 61 (list); Attems 1953: 155

(list); Jeekel 1971: 27 (list); Jeekel 1974: 46 (classifica¬

tion); Hoffman 1976: 119 (classification); Hoffman 1980:

63 (list); Jeekel 2001: 16 (list); Mauries 2001: 131 (clas¬

sification); Wesener & VandenSpiegel 2009: 551 (list).

Synonyms:

— Chinosphaera Attems, 1935: 130, synonymized by

Mauries 2001: 139. Attems 1936: 153; Attems 1943: 61;

Jeekel 1971: 24; Jeekel 1974: 46; Hoffman 1976: 119;

Hoffman 1980: 63; Jeekel 2001: 13; Mauries 2001: 132.

Tvpe-species : Chinosphaera maculosa Attems.

Description: Anterior telopods similar to those of Cryxus ,

usually consisting of three telopoditomeres. Telopodito-

mere 2 posteriorly with a ledge-like process. Telopodito-

mere 3 apically completely bent posteriorly and sharp-

edged, forming a clamp with posterior process of

telopoditomere 2. In some species a rudimentary telopodit-

omere 4 is present. Posterior telopods with four telopodit¬

omeres, telopoditomere 2 elongated. Telopoditomere 2

apically on posterior side, often with a sclerotized knob

towards telopoditomeres 3 and 4 with a single or bilobed

membranous process. Tarsus with a single apical spine.

Members 20-35 mm long.

Type-species: Prionobelum durum Verhoeff.

Species included (7)

Prionobelum durum Verhoeff, 1924 (Vietnam)

Prionobelum hainani (Gressitt, 1941) (China: Hainan)

Prionobelum joliveti Mauries, 2001 (China: Hainan)

Prionobelum leve (Attems, 1938) (Vietnam)

Prionobelum maculosum (Attems, 1935) (China: Fukien)

Prionobelum majorinum (Zhang & Lie, 1982) (China:

Hainan)

Prionobelum multidentata (Wang & Zhang, 1993) (Chi¬

na: Fujian)

Prionobelum durum Verhoeff, 1924

Prionobelum durum Verhoeff, 1924: 64 (first description)

Prionobelum durum.— Attems 1936: 192 (list)

Prionobelum durum — Moritz & Fischer 1974: 368 (list)

Prionobelum durum, var. nitidum — Verhoeff 1924: 65

(new locality)

Prionobelum durum — Golovatch 1983 (list)

Prionobelum durum — Jeekel 2001: 16 (list)

Prionobelum durum nitidum — Mauries 2001: 132 (treat¬

ed as subspecies instead of variety)

Prionobelum durum durum — Enghoff et al. 2004: 32 (list)

Prionobelum durum nitidum :— Enghoff et al. 2004: 32

(list).

Remarks: Variety P. durum var. nitidum comes from the

same locality as the type of P. durum. The varieties were

treated as subspecies by Mauries (2001: 132) and Enghoff

et al. (2004: 32), a move not followed here.

Types: M syntype.

Habitat: Vietnam: Tonkin.

Deposited: ZMB 5745 (Moritz & Fischer 1974: 368).

Prionobelum hainani (Gressit, 1941)

Zephronia hainani Gressit, 1941: 56 (first description)

Zephronia hainani — Mauries 2001: 134 (redescription)

Prionobelum hainani :— Mauries 2001: 139 (new comb.).

Types: Holotype, paratypes.

Habitat: Tai-Pin-ts’uen (Dwa Bi), 375 m, foot of Loi

Mother Mountain (19°N, 109°40’E), Hainan, China.

Deposited: CAS (Mauries 2001: 134), Lingnan Universi¬

ty, Canton, China (Gressitt 1941: 55).

Prionobelum joliveti Mauries, 2001

Prionobelum joliveti Mauries, 2001: 136 (first descrip.).

Types: M holotype, 1 F allotype, 12 M, 4 F paratypes

Habitat: China, Hainan, ‘sur une colline a louest de la ville

nouvelle (et universitaire) de Danzhou (109°33’E,

19°30’N), dans une plantation d’heveas’.

Deposited: Holotype and paratypes MNHN CB054, 1 M,

1 F paratypes CAS, 1 M, 1 F paratypes ZMUM.

Prionobelum leve (Attems, 1938)

Chinosphaera levis Attems, 1938: 198 (first description)

Chinosphaera levis — Golovatch 1983 (list)

Chinosphaera levis — Jeekel 2001: 14 (list)

Prionobelum levis [sic]:— Mauries 2001: 139 (new com¬

bination)

Prionobelum leve — Enghoff et al. 2004: 32 (list).

Types: M and F syntypes.

Habitat: Vietnam: Cauda, Nhalrang, South Annam; Nin-

hoa, Nhalrang; Groa, Provo Khan Hoa, South Annam.

Deposited: NHMW 2197 (Stagl & Schiller, in prep.).

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Thomas Wesener

Prionobelum maculosum (Attems, 1935)

Chinosphaera maculosa Attems, 1935: 134 (first descrip.)

Chinosphaeramaculosa.— Attems 1936: 191 (list)

Chinosphaera maculosa -- Attems 1938: 200 (list)

Chinosphaera maculosa.— Jeekel 2001: 14 (list)

Chinosphaera maculosa.— Mauries 2001: 134 (redescrip¬

tion)

Prionobelum maculosum :-- Mauries 2001: 139 (new com¬

bination).

Types: M holotype.

Habitat: China: Foochow, Prov. Fokien.

Deposited: ZMUH (Mauries 2001: 134).

Prionobelum majorinum (Zhang & Lie, 1982)

Chinosphaera majorina Zhang & Li, 1982: (first descrip.)

Prionobelum majorinum :— Mauries 2001: 139 (new com¬

bination).

Types: M holotype and 1 M and 4 F paratypes.

Habitat: China, Hainan, S. of the island, Diaoluoshan,

18,47°N, 109,52°E.

Deposited: Institute of Zoology, Academia Sinica, holo¬

type 078058-1, F paratype 078058-2.

Prionobelum multidentata (Wang & Zhang, 1993)

Chinosphaera multidentata Wang & Zhang, 1993: 849

(first description)

Prionobelum multidentata — Mauries 2001: 139 (new

combination).

Types: M holotype.

Habitat: China, Fujian, Mount Longqi.

Deposited: Institute of Zoology, Academia Sinica, 910817-

1 .

Leptoprotopus Silvestri. 1897

Leptoprotopus Silvestri, 1897c: 645 (first description);

Jeekel 1971: 26 (list); Hoffman 1976: 119 (list); Hoffman

1980: 63 (list); Jeekel 1974: 46 (list, as synonym of

Sphaeropoeus ); Jeekel 2001: 22 (list); Mauries 2001: 140

(list); Wesener & VandenSpiegel 2009: 551 (list).

Description: First two male legs with greatly enlarged

claws, claw as long as tarsus, a unique character for the

order. Tooth of mandible also enormously enlarged and

elongated. Anterior and posterior telopods greatly resem¬

bling those of Sphaeropoeus. 15 mm long.

Type-species: Sphaeropoeus gladiator Pocock, 1894.

Monotypic

Remarks: This genus was listed as a synonym of

Sphaeropoeus (Jeekel 1974, Mauries 2001). Because no

argument for this synonymization was given, and the char¬

acters of Sphaeropoeus gladiator are unique for the

Sphaerotheriida, Leptoprotopus is kept separate from

Sphaeropoeus.

Leptoprotopus gladiator (Pocock, 1894)

Sphaeropoeus gladiator Pocock, 1894: 327 (first descrip.)

Sphaeropoeus gladiator— Silvestri 1895: 723 (list)

Sphaeropoeus gladiator :— Pocock 1895: 412 (list)

Leptoprotopus gladiator— Silvestri 1897: 645 (new com¬

bination)

Sphaeropoeus gladiator— Attems 1914: 144 (list)

Leptoprotopus gladiator— Jeekel 2001: 22 (list).

Types: M holotype.

Habitat: Indonesia: Sumatra, Paningahan.

Deposited: NHML (Online Catalogue).

Sphaeropoeus Brandt. 1833

Sphaeropoeus Brandt, 1833: 199 (first description);

Brandt 1841a: 364 (description); Gervais 1847: 83 (list);

C.L. Koch 1847: 38 (description); Humbert 1865: 37 (new

species); Pocock 1894: 326 (new species); Pocock 1895:

412 (new species); Silvestri 1896b: 205; Flower 1901: 7

(new species); Attems 1914: 142 (list); Verhoeff 1924: 59

(list); Attems 1926: 120 (list); Attems 1928: 203 (list); At¬

tems 1935: 130 (list) ; Attems 1936: 152 (list); Attems

1943: 61 (list); Jeekel 1951: 105 (new species); Attems

1953: 155 (list); Jeekel 1971: 28 (list); Jeekel 1974: 46

(classification); Hoffman 1976: 119 (list); Hoffman 1980:

63 (list); Hugot et al. 1991: 65 (parasites); Jeekel 2001:

22 (list); Mauries 2001: 140 (list); Wesener & Vanden¬

Spiegel 2009: 551 (morphological phylogenetic analysis).

Synonyms

— Tonkinobelum Verhoeff, 1924: 58 (first description),

synonymized by Jeekel 1974: 46.

Attems 1926: 120 (list); Attems 1928: 203 (list); Verho¬

eff 1929: 1384 (description); Attems 1935: 128 (list);

Attems 1936: 152 (list); Attems 1943: 62 (list); Jeekel

1971: 29 (list); Hoffman 1976: 119 (list); Hoffman 1980:

63 (list).

Type species : Tonkinobelum maculatum Verhoeff, 1924.

— Pantitherium Attems, 1932: 7 (first description), syn¬

onymized by Jeekel 1974: 46. Attems 1935: 131 (list);

Attems 1943: 63 (list); Attems 1953: 155 (list); Jeekel

1971:27 (list).

Type species : Pantitherium lugubre Attems, 1932.

— Lissosphaera Attems, 1943: 62 (first description), syn¬

onymized by Jeekel 1974: 46. Jeekel 1971: 26 (list).

Type species : Lissosphaera speciosa Attems, 1943.

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Species Catalogue Sphaerotheriida

Description: Anterior telopod with four podomeres dis¬

tal to syncoxite. Telopoditomeres 2 and 3 each almost as

long as telopoditomere 1. A specially shaped lateral

process of telopoditomere 2 and a process of telopodito¬

mere 3 form a clamp. Posterior telopods with four

telopoditomeres, immovable finger slender, with membra¬

nous process. Telopoditomere 3 with small sclerotized

teeth and white ledge. Telopoditomere 4 lacking teeth or

spines, with a mambranous ledge. 20-50 mm long. Api¬

cal antennomere axe-shaped, with up to 180 apical cones

(record number of the order). Tarsi with a single apical

spine, rarely two. In urgent need of a revision. The status

of many species assigned to this genus is unclear.

Type-species: Sphaeropoeus Hercules Brandt, 1833.

Species included (20)

Sphaeropoeus extinctus Silvestri, 1895 (Malaysia, Indone¬

sia: Nias)

Sphaeropoeus gisleni (Verhoeff, 1939) (Sumatra)

Sphaeropoeus Hercules Brandt, 1833 (Sumatra, Malaysia)

Sphaeropoeus lugubris (Attems, 1932) (Sumatra)

Sphaeropoeus maculatus Verhoeff, 1924 (Vietnam)

Sphaeropoeus magnus Chamberlin, 1921 (Borneo:

Sarawak)

Sphaeropoeus malaccanus Verhoeff, 1910-1914 (Singa¬

pore)

Sphaeropoeus modiglianii Silvestri, 1895 (Sumatra,

Malay Peninsula)

Sphaeropoeuspellitus (Attems, 1943) (Sumatra)

Sphaeropoeuspolitus (Attems, 1935) (Sumatra)

Sphaeropoeuspunctulatissimus Silvestri, 1897 (Sumatra)

Sphaeropoeus speciosus (Attems, 1943) (Sumatra)

Sphaeropoeus stollii Pocock, 1895 (Java)

Sphaeropoeus sumatrensis (Chamberlin, 1945) (Sumatra)

Sphaeropoeus tatusiaeformis Daday, 1889 (Sumatra)

Sphaeropoeus tigratus Silvestri, 1897 (Sumatra)

Sphaeropoeus tuberculosus Karsch, 1881 (Borneo)

Sphaeropoeus unciger (Attems, 1943) (Sumatra)

Sphaeropoeus variegatus Pocock, 1895 (unknown)

Sphaeropoeus velutinus Carl, 1906 (Sumatra)

Sphaeropoeus extinctus Silvestri, 1895

Sphaeropoeus extinctus Silvestri, 1895: 722 (first descrip.)

Sphaeropoeus extinctus :— Sinclair 1901: 527 (description)

Sphaeropoeus extinctus -- Attems 1914: 143 (list)

Sphaeropoeus extinctus.— Jeekel 2001: 22 (list).

Remark: The spelling “exstinctus” at the heading of the

original description is an obvious printer’s error (Jeekel

2001 ).

Types: M and F syntypes.

Habitat: Indonesia: Nias, Hili Zabobo. Malaysia:

Tapelung, Patalung State.

Deposited: IEPN?.

Sphaeropoeus gisleni (Verhoeff, 1939)

Tonkinobelum gisleni Verhoeff, 1939: 2 (first description)

Tonkinobelum gisleni :— Wang 1967: 484 (list)

Sphaeropoeus gisleni :— Jeekel 2001: 23 (list, new comb.).

Types: M holotype.

Habitat: Indonesia: Sumatra.

Deposited: MZLU.

Sphaeropoeus glabrus (Attems, 1943)

Arthrosphaera glabra Attems, 1943: 72 (first description)

Arthrosphaera glabra:— Wang 1967: 484 (list)

Sphaeropoeus glabrus :— Jeekel 2001: 23 (list, new comb.).

Types: 2 M syntypes.

Habitat: Indonesia: Sumatra, Padang.

Deposited: NHMW2215.

Sphaeropoeus hercules Brandt, 1833

Sphaeropoeus hercules Brandt, 1833: 200 (first descrip.)

Zephronia Hercules — Gervais 1837: 43 (list)

Sphaeropoeus Hercules — Brandt 1841a: 364 (new local.)

Zephronia Hercules — Gervais 1847: 84 (list)

Sphaeropoeus hercules :— CL. Koch 1863: 3 (description)

Sphaeropoeus Hercules — Karsch 1881: 23 (description)

Zephronia hercules — Preudhomme de Borre 1884: 24

(list)

Sphaeropoeus Hercules — Daday 1889: 141 (new local.)

Sphaeropoeus Hercules — Pocock 1894: 326 (description)

Sphaeropoeus Hercules — Silvestri 1895: 722 (new local.)

Sphaeropoeus Hercules — Pocock 1895: 412 (new local.)

Sphaeropoeus hercules :— Sinclair 1901: 527 (new local.)

Sphaeropoeus hercules — Carl 1906: 230 (description)

Sphaeropoeus Hercules — Attems 1914: 143 (list)

Sphaeropoeus hercules :— Moritz & Fischer 1973: 369

(list)

Sphaeropoeus Hercules - Hoffman 1980: 62 (list)

Sphaeropoeus Hercules — Jeekel 2001: 23 (list)

Sphaeropoeus Hercules — De Zoysa et al. 2016: 456 (list)

— Sphaeropoeus banksianus Butler, 1873: 181 (first de¬

scrip.), synonymized under S. hercules by Pocock, 1894:

326

Zephronia Banksiana [sic!]:— Preudhomme de Borre

1884: 23 (list)

— Zephronia barbata Butler, 1882: 197 (first descrip.),

synonymized under S. hercules by Pocock 1894: 326

Zephronia barbata :— Preudhomme de Borre 1884: 23

(list)

Zephronia barbarta [sic!]:— Wang 1967: 484 (list).

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Remark: De Zoysa et al. (2016) list S. Hercules from Sri

Lanka, citing the wrong record by Pocock (1892) thatwas

later corrected by the author.

Types: M holotype.

Habitat: Unknown (Type). Indonesia: Sumatra, Padang,

Manindjau, Singkarah, Paningahan, Ajer Mantjur; Palem-

bang; Tibing Tingie; M. Singalang, Ajer Mantdor, Kaju

Tanam; Si Rambe; Palembang; Malaysia.

Deposited: M holotype, ZMB 54 (Moritz & Fischer 1973:

369).

Sphaeropoeus lugubris (Attems, 1932)

Pantitherium lugubre Attems, 1932: 8 (first description)

Panitherium [sic!] lugurbe [sic!]:-Wang 1967: 484 (list).

Sphaeropoeus lugubris.— Jeekel 2001: 23 (list, new com¬

bination).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Panti.

Deposited: NHMW 3955 (Stagl & Schiller, in prep.).

Sphaeropoeus maculatus (Verhoeff, 1924)

Tonkinobelum maculatum Verhoeff, 1924: 62 (first de¬

scription)

Tonkinobelum maculatum.— Attems 1936: 192 (list)

Tonkinobelum maculatum — Wang 1967: 484 (list)

Tonkinobelum maculatum :— Korsos 1983: 118 (list)

Tonkinobelum maculatum :— Golovatch 1983 (list)

Sphaeropoeus maculatus — Jeekel 2001: 23 (list)

Sphaeropoeus maculatus — Enghoff et al. 2004: 32 (list).

Remark: According to Jeekel (2001), this species might

represent a synonym of S. tigratus Silvestri, which got the

wrong label.

Types: M syntype.

Habitat: Vietnam: Tonkin, Man Son.

Depository: HNMB 2858/1 (Korsos 1983).

Sphaeropoeus magnus Chamberlin, 1921

Sphaeropoeus magnus Chamberlin, 1921: 62 (first de¬

scription)

Sphaeropoeus magnus :— Wang 1967: 484 (list)

Sphaeropoeus magnus :— Jeekel 2001: 23 (list).

Types: M holotype.

Habitat: Sarawak: Ladong.

Deposited: MCZ 4708.

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Sphaeropoeus malaccanus Verhoeff, 1910

Sphaeropoeus malaccanus Verhoeff 1910-1914: pi. 5 fig.

77 (first description)

Sphaeropoeus malaccanus :— Verhoeff 1924: 68 (descrip.)

Sphaeropoeus malaccanus — Jeekel 2001: 23 (list)

Sphaeropoeus malaccanus — Decker 2013: 3 (list).

Types: 3 M, 1 F syntypes.

Habitat: Singapore.

Deposited: Unknown.

Sphaeropoeus modiglianii Silvestri, 1895

Sphaeropoeus modiglianii Silvestri, 1895: 722 (first de¬

scription)

Sphaeropoeus modiglianii — Sinclair 1901: 527 (new lo¬

cality)

Sphaeropoeus modiglianii :— Attems 1914: 143 (list)

Sphaeropoeus modiglianii :— Jeekel 2001: 23 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Si Rambe (Type). “Malay

Peninsula”.

Deposited: Unknown.

Sphaeropoeus pellitus (Attems, 1943)

Kophosphaera pellita Attems, 1943: 66 (first description)

Kophosphaerapellita :— Wang 1967: 485 (list)

Sphaeropoeus pellitus :— Jeekel 2001: 24 (list, new com¬

bination).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra.

Deposited: NHMW 2194.

Sphaeropoeuspolitus (Attems, 1935)

Pantitherium politum Attems, 1935: 121 (first description)

Panitherium [sic!] politum — Wang 1967: 484 (list)

Sphaeropoeus politus :— Jeekel 2001: 24 (list, new com¬

bination).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Singkarak, Kepajang-

Benkulen.

Deposited: NHMW 2217, 2218 (Stagl & Schiller, in

prep.).

Sphaeropoeuspunctulatissimus Silvestri, 1897

Sphaeropoeuspunctulatissimus Silvestri, 1897a: 14 (first

description)

Sphaeropoeus punctulatissimus :— Attems, 1914: 143 (list)

Sphaeropoeus punctulatissimus :— Jeekel 2001: 24 (list).

©ZFMK

Species Catalogue Sphaerotheriida

Types: F syntypes.

Habitat: Indonesia: Sumatra, Deli.

Deposited: MTD 280.

Sphaeropoeus speciosus (Attems, 1943)

Lissosphaera speciosa Attems, 1943: 63 (first description)

Lissosphaera speciosa.— Wang 1967: 485 (list)

Sphaeropoeus speciosus :— Jeekel 2001: 24 (list, new com¬

bination).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra.

Deposited: NHMW 2219 (Stagl & Schiller, in prep.).

Sphaeropoeus stollii Pocock, 1895

Sphaeropoeus stollii Pocock, 1895: 412 (first description)

Sphaeropoeus stollii.— Attems 1914: 143 (list)

Sphaeropoeus stollii — Jeekel 2001: 24 (list).

Types: Unknown.

Habitat: Indonesia: Java: Djakarta.

Deposited: NHML?

Sphaeropoeus sumatrensis (Chamberlin, 1945)

Tonkinobelum sumatrense Chamberlin, 1945: 2 (first de¬

scription)

Tonkinobelum sumatrensis [sic!]:—Wang 1967: 485 (list)

Sphaeropoeus sumatrensis :— Jeekel 2001: 24 (list, new

combination).

Types: 2 M syntypes.

Habitat: Indonesia: Sumatra, Manindjau.

Deposited: AMNH.

Sphaeropoeus tatusiaeformis Daday, 1889

Sphaeropoeus tatusiaeformis Daday, 1889: 141 (first de¬

scription)

Sphaeropoeus tatusiaeformis :— Attems 1914: 144 (list)

Sphaeropoeus tatusiaeformis :— Wang 1967: 384 (list)

Sphaeropoeus tatusiaeformis — Korsos 1983: 119 (list)

Sphaeropoeus tatusiaeformis :— Jeekel 2001: 24 (list).

Types: Syntypes.

Habitat: Indonesia: Sumatra.

Depository: HNMB 648.14.d/5.

Sphaeropoeus tigratus Silvestri, 1897

Sphaeropoeus tigratus Silvestri, 1897a: 14 (first descrip¬

tion)

Sphaeropoeus tigratus :— Carl 1922: 565 (new locality)

Sphaeropoeus tigratus :— Attems 1914: 143 (list)

Sphaeropoeus tigratus :— Jeekel 2001: 24 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Deli; Sumatra Bandar

Baroe, Deli.

Deposited: MTD 278.

Sphaeropoeus tuberculosus Karsch, 1881

Sphaeropoeus tuberculosus Karsch, 1881: 33 (first de¬

scription)

Zephronia tuberculosa :— Preudhomme de Borre 1884: 25

(list)

Sphaeropoeus tuberculosus — Attems 1914: 144 (list)

Sphaeropoeus tuberculosus :— Moritz & Fischer, 1974: 370

(list)

Sphaeropoeus tuberculosus — Jeekel 2001: 24 (list).

Types: F holotype (Moritz & Fischer 1974: 370).

Habitat: Indonesia: Kalimantan, Benkajang.

Deposited: ZMB 590.

Sphaeropoeus unciger { Attems, 1943)

Kophosphaera unciger a Attems, 1943: 66 (first descrip.)

Sphaeropoeus unciger— Jeekel 2001: 24 (new combina¬

tion, list).

Types: M holotype.

Habitat: Indonesia: Sumatra.

Deposited: NHMW 2195.

Sphaeropoeus variegatus Pocock, 1895

Sphaeropoeus variegatus Pocock, 1895: 412 (first descrip¬

tion)

Sphaeropoeus variegatus — Jeekel 2001: 24 (list).

Types: M holotype, .

Habitat: “?, Near Singapore”.

Deposited: NHML.

Sphaeropoeus velutinus Carl, 1906

Sphaeropoeus velutinus Carl, 1906: 232 (first description)

Sphaeropoeus velutinus :— Attems 1914: 144 (list)

Sphaeropoeus velutinus :— Jeekel 2001: 24 (list)

— Sphaeropoeus velutinus, var. xanthopleurus Carl,

1909: 249 (first description)

Sphaeropoeus velutinus, var. xanthopleurus :— Attems,

1914: 144 (list)

Sphaeropoeus velutinus, var. xanthopleurus :— Jeekel

2001: 25 (list).

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Thomas Wesener

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Deli; Karoo plain at foot of

Mt. Si-Nabung.

Deposited: MHNG.

Tigridosphaera Jeekel. 2000

Tigridosphaera Jeekel, 2000: 68 (first description); Jeekel

2001: 25 (list); Wesener & VandenSpiegel 2009: 551 (list).

Description: Anterior telopods with three podomeres dis¬

tal to syncoxite. Telopoditomere 3 bent posteriad, form¬

ing clamps with the process of telopoditomere 2. Poste¬

rior telopods with four telopoditomeres. Tarsi with a sin¬

gle apical spine. Likely synonymous with Prionobelum.

Type-species: Sphaeropoeusglobusmagicus Jeekel, 1951.

Species included (4)

Tigridosphaera bimaculata (Pocock, 1895) (Singapore)

Tigridosphaera evansi (Sinclair, 1901) (Malay Peninsu¬

la)

Tigridosphaera globusmagicus (Jeekel, 1951) (Malaysia)

Tigridosphaera zonata (Pocock, 1895) (Malaysia)

Tigridosphaera bimaculata (Pocock, 1895)

Sphaeropoeus bimaculatus Pocock, 1895: 412 (first de¬

scription)

Sphaeropoeus bimaculatus -- Attems 1914: 143 (list)

Sphaeropoeus bimaculatus.— Wang 1967: 484 (list)

Tigridosphaera bimaculata :— Jeekel 2001: 25 (new com¬

bination)

Tigridosphaera bimaculata :— Decker 2013: 3 (list).

Types: Unknown

Habitat: Singapore.

Deposited: NHML?

Tigridosphaera evansi (Sinclair, 1901)

Sphaeropoeus evansi Sinclair, 1901: 526 (first description)

Sphaeropoeus evansi :— Attems 1914: 143 (list)

Tigridosphaera evansi :— Jeekel 2000: 69 (new comb.)

Tigridosphaera evansi — Jeekel 2001: 25 (list)

Tigridosphaera evansi — Decker 2013: 3 (list).

Types: M and F syntypes.

Habitat: “Malay Peninsula”.

Deposited: NHML?

Tigridosphaera globusmagicus (Jeekel, 1951)

Sphaeropoeus globus-magicus Jeekel, 1951: 101 (first de¬

scription)

Tigridosphaera globusmagicus :— Jeekel 2000: 68 (new

combination)

Tigridosphaera globusmagicus :— Jeekel 2001: 25 (list).

Types: M holotype, 1 M paratype.

Habitat: Malaysia: Kepong, Bukit Lebong Forest Reserve,

Selangor.

Deposited: Holotype ZMA, paratype Raffles Museum Sin¬

gapore.

Tigridosphaera zonata (Pocock, 1895)

Sphaeropoeus zonatus Pocock, 1895: 412 (first descrip¬

tion)

Sphaeropoeus zonatus — Attems 1914: 144 (list)

Sphaeropoeus zonatus — Wang 1967: 484 (list)

Tigridosphaera zonata — Jeekel 2001: 25 (list, new com¬

bination).

Types: Holotype of unknown sex.

Habitat: Malaysia: Malacca.

Deposited: NHML (Online Catalogue).

Subfamily Castanotheriinae Jeekel

Jeekel, 1974: 46; Castanotheriini Hoffman, 1976: 119;

Hoffman, 1980: 63; Jeekel 2001: 25; Mauries 2001: 141.

Castanotherium Pocock. 1895

(Figs 1 IB, C)

Castanotherium Pocock, 1895: 413 (first description); At¬

tems 1914: 142 list); Verhoeff 1924: 58 (list); Attems

1926: 120 (list); Attems 1928: 203 (list); Attems 1935: 132

(list); Attems 1943: 62 (list); Attems 1953: 155 (list);

Jeekel 1971: 23 (list); Jeekel 1974: 46 (classification);

Hoffman 1976: 120 (classification); Hoffman 1980: 64

(list); Hugot et al. 1991: 65 (parasites); Mauries 2001: 141

(list); Wesener & VandenSpiegel 2009: 551 (morpholog¬

ical phylogenetic analysis).

Synonyms:

— Borneopoeus Verhoeff, 1924: 57, synonymized with

Castanotherium by Hoffman 1976: 120. Attems 1926:

120; Attems 1928: 203; Verhoeff 1929: 1383; Attems

1935: 127; Attems 1936: 152; Attems 1943: 62; Attems

1953: 155; Jeekel 1971: 23; Jeekel 1974: 46; Jeekel

2001: 25.

Tvpe-species : Borneopoeus costatus Verhoeff, 1924.

— Pulusphaera Attems, 1935: 129, synonymized with

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Species Catalogue Sphaerotheriida

Castanotherium by Hoffman 1976: 120. Attems 1943:

62; Attems 1953: 155; Jeekel 1971: 28; Jeekel 1974: 46.

Tvpe-species : Pulusphaera fern Attems, 1935.

Description: Posterior and anterior telopods with only

three podomeres distal to syncoxite. Immovable finger of

posterior telopod greatly enlarged, in some species twice

as wide as podomere 3. Antennae axe-shaped, with nu¬

merous apical cones. Thoracic shield at margin to groove

often with ridges. Tarsi with a single apical spine.

Type-species: Castanotherium hosei Pocock, 1895.

Species included (50)

Castanotherium amythrum (Attems, 1897) (Indonesia:

Halmahera)

Castanotherium boetonense Carl, 1912 (Indonesia: Bu-

tung)

Castanotherium carinatum (Pocock, 1890) (Borneo)

Castanotherium castaneum (Newport, 1844) (Philippines)

Castanotherium celebense Silvestri, 1897 (Indonesia: Su¬

lawesi)

Castanotherium cinctum (Carl, 1906) (Sumatra)

Castanotherium conspicuum Silvestri, 1897 (Borneo)

Castanotherium costatum (Verhoeff, 1924) (Borneo)

Castanotherium criniceps (Attems, 1897) (Indonesia: Su¬

lawesi)

Castanotherium crinitum (Attems, 1935) (Borneo)

Castanotherium decoratum Carl, 1912 (Indonesia: Su¬

lawesi)

Castanotherium distinctum Carl, 1912 (Indonesia: Sulawe¬

si)

Castanotherium dorsispina (Attems, 1936) (Borneo)

Castanotherium everettii Pocock, 1895 (Borneo)

Castanotherium extraneum (Attems, 1943) (Java)

Castanotherium falcicorne (Tomosvary, 1885) (Borneo)

Castanotherium ferum (Attems, 1935) (Borneo, Bismar¬

ck Islands?)

Castanotherium fulvicorne Pocock, 1895 (Philippines)

Castanotherium glabratum (Newport, 1844) (Philip¬

pines, Borneo?)

Castanotherium granulatum (Tomosvary, 1885) (Borneo)

Castanotherium hirsutellum Pocock, 1895 (Philippines)

Castanotherium hosei Pocock, 1895 (Borneo)

Castanotherium humile (Silvestri, 1895) (Indonesia: Bua-

Bua)

Castanotherium ignobile (Butler, 1872) (Java)

Castanotherium indigenum (Attems, 1943) (Borneo)

Castanotherium innominatum (Butler, 1872) (Philip¬

pines)

Castanotherium insigne (Brandt, 1833) (Java)

Castanotherium laeve Carl, 1912 (Indonesia: Sulawesi)

Castanotherium lamprongse (Chamberlin, 1945) (Suma¬

tra)

Castanotherium leium Chamberlin, 1921 (Philippines)

Castanotherium moderatum (Chamberlin, 1945) (Java)

Castanotherium nigriceps (Pocock, 1894) (Java)

Castanotherium nigromaculatum Silvestri, 1897 (Borneo)

Castanotherium obscurum Chamberlin, 1921 (Borneo)

Castanotherium ornatum Carl, 1912 (Indonesia: Sulawe¬

si)

Castanotherium pellitum Attems, 1932 (Borneo)

Castanotherium pilosum Carl, 1912 (Indonesia: Sulawe¬

si)

Castanotheriumplasoni (Attems, 1943) (Indonesia: Java)

Castanotherium pyrrhomelanum (Attems, 1897) (Borneo)

Castanotherium rufipes (Pocock, 1894) (Indonesia: Java)

Castanotherium simplex (Carl, 1909) (Indonesia: Java)

Castanotherium sparsepunctatum Carl, 1912 (Indonesia:

Sulawesi)

Castanotherium stellatum Carl, 1912 (Indonesia: Sulawe¬

si)

Castanotherium sulcicollis (Karsch, 1881) (Java, Borneo,

Philippines?)

Castanotherium suspectum Carl, 1912 (Indonesia: Su¬

lawesi)

Castanotherium tivium (Chamberlin, 1945) (Indonesia:

Sumatra)

Castanotherium tricollis (Karsch, 1881) (Indonesia:

Sumatra, Java?)

Castanotherium variegatum (Attems, 1943) (Borneo)

Castanotherium volzi (Carl, 1906) (Indonesia: Sumatra)

Castanotherium whiteheadi Pocock, 1895 (Borneo)

Castanotherium amythrum (Attems, 1897)

Zephronia amythra Attems, 1897: 483 (first description)

Zephronia amythra.— Attems 1914: 144 (list)

Castanotherium amythrum :— Chamberlin 1920: 91 (list)

Castanotherium amythrum :— Jeekel 2001: 26 (list).

Types: F holotype.

Habitat.: Indonesia: Halmahera, Soah Konorah.

Deposited: NHMW?

Castanotherium boetonense Carl, 1912

Castanotherium boetonense Carl, 1912: 119 (first descrip.)

Castanotherium boetonense :— Attems 1914: 149 (list)

Castanotherium boetonense :— Chamberlin 1920: 91 (list)

Castanotherium boetonense :— Jeekel 2001: 26 (list).

Types: F holotype.

Habitat: Indonesia: Butung island, SE of Sulawesi.

Deposited: MHNG.

Castanotherium carinatum (Pocock, 1890)

Zephronia carinata Pocock, 1890a: 82 (first description)

Zephronia carinata — Attems 1914: 144 (list)

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Castanotherium carinatum — Chamberlin 1921: 69 (list)

Castanotherium carinatum.— Jeekel 2001: 26 (list).

Type: 1 M, 1 F syntypes.

Habitat: North Borneo: Mt. Kina Balu.

Deposited: NHML (Online Catalogue).

Castanotherium castaneum (Newport, 1844)

Zephronia castanea Newport, 1844: 265 (first description)

Zephronia castanea :-- Butler 1873: 181 (list)

Zephronia castanea — Preudhomme de Borre 1884: 24

(list)

Zephronia castanea :-- Attems 1914: 145 (list)

Castanotherium castaneum — Jeekel 2001: 26 (list, new

combination).

Types: Holotype of unknown sex.

Habitat: Philippines.

Deposited: NHML (Online Catalogue).

Castanotherium celebense Silvestri, 1897

Castanotherium celebense Silvestri, 1897a: 15 (first de¬

scription)

Castanotherium celebense — Carl 1912: 86 (list)

Castanotherium celebense — Attems 1914: 149 (list)

Castanotherium celebense — Chamberlin 1920: 90 (list)

Castanotherium celebense — Jeekel 2001: 26 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sulawesi, Minahassa.

Deposited: MTD 282.

Castanotherium cinctum (Carl, 1906)

Sphaeropoeus cinctus Carl, 1906: 235 (first description)

Castanotherium cinctum — Attems 1914: 149 (list)

Castanotherium cinctum — Jeekel 2001: 26 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra.

Deposited: MHNG.

Castanotherium conspicuum Silvestri, 1897

Castanotherium conspicuum Silvestri, 1897a: I (first de¬

scription)

Castanotherium conspicuum :— Attems 1914: 149 (list)

Castanotherium conspicuum :— Jeekel 2001: 26 (list).

Types: F holotype.

Habitat: Indonesia: Southwest Kalimantan.

Deposited: MTD 282.

Castanotherium costatum (Verhoeff, 1924)

Borneopoeus costatus Verhoeff, 1924: 60 (first descrip-

ciption)

Castanotherium costatum :— Jeekel 2001: 26 (list, new

combination).

Types: 3 M, 1 F syntypes.

Habitat: “Central Borneo”.

Deposited: NHML (Online Catalogue).

Castanotherium criniceps (Attems, 1897)

Zephronia criniceps Attems, 1897: 482 (first description)

Castanotherium criniceps :— Carl 1906: 234 (list)

Castanotherium criniceps :— Carl 1912: 86 (description)

Castanotherium criniceps :— Attems 1914: 149 (list)

Castanotherium criniceps :— Chamberlin 1920: 90 (list)

Castanotherium criniceps :— Jeekel 2001: 26 (list).

Types: M holotype.

Habitat: Indonesia: Sulawesi, Minahassa (Type); North

Sulawesi: Vokano Soputan, Tomohon.

Deposited: NHMW?

Castanotherium crinitum (Attems, 1935)

Pulusphaera crinita Attems, 1935: 138 (first description)

Castanotherium crinitum :— Jeekel 2001: 26 (list, new

combination).

Types: M holotype.

Habitat: Indonesia: Kalimantan, Bt. Abat.

Deposited: NHMW?

Castanotherium decoratum Carl, 1912

Castanotherium decoratum Carl, 1912: 118 (first descrip¬

tion)

Castanotherium decoratum — Attems 1914: 150 (list)

Castanotherium decoratum :— Chamberlin 1920: 91 (list)

Castanotherium decoratum :— Jeekel 2001: 26 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sulawesi, Loka, near Bonthain, South

Sulawesi.

Deposited: MHNG.

Castanotherium distinctum Carl, 1912

Castanotherium distinctum Carl, 1912: 106 (first descrip¬

tion)

Castanotherium distinctum — Attems 1914: 150 (list)

Castanotherium distinctum :— Chamberlin 1920: 90 (list)

Castanotherium distinctum :— Jeekel 2001: 27 (list).

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Species Catalogue Sphaerotheriida

Types: M holotype.

Habitat: Indonesia: Sulawesi, Ussu (Southeast Sulawesi).

Deposited: MHNG.

Castanotherium dorsispina (Attems, 1936)

Borneopoeus dorsispina Attems, 1936: 193 (first descrip.)

Castanotherium dorsispina:— Jeekel 2001: 27 (list, new

combination).

Types: F holotype.

Habitat: Sarawak: Kapit.

Deposited: NHMW, unclear if types (Stagl & Schiller, in

prep.).

Castanotherium everettii Pocock, 1895

Castanotherium everettii Pocock, 1895: 414 (first descrip.)

Castanotherium everettii :-- Attems 1914: 150 (list)

Castanotherium everettii.— Jeekel 2001: 27 (list).

Type: Syntypes?

Habitat: Sarawak.

Deposited: NHML (Online Catalogue).

Castanotherium extraneum (Attems, 1943)

Pulusphaera extranea Attems, 1943: 70 (first description)

Castanotherium extraneum :-- Jeekel 2001: 27 (list, new

combination).

Types: M holotype.

Habitat: Indonesia: Java, Samarang.

Deposited: NHMW 118/70 (Stagl & Schiller, in prep.).

Castanotherium falcicorne (Tomosvary, 1885)

Sphaeropoeus falcicornis Tomosvary, 1885: 68 (first de¬

scription)

Sphaeropoeus falcicornis — Silvestri 1896a: 21 (list)

Sphaeropoeus falcicornis — Attems 1914: 144 (list)

Sphaeropoeus falcicornis — Korsos 1983: 118 (list)

Castanotherium falcicorne :-- Jeekel 2001: 27 (list, new

combination).

Types: Syntypes.

Locality: Indonesia, Kalimantan, Matang.

Deposited: HNHM 305.62/17 (Korsos 1983).

Castanotherium ferum (Attems, 1935)

Pulusphaera fera Attems, 1935: 136 (first description)

Pulusphaera fera — Attems 1943:71 (new locality)

Castanotherium ferum — Jeekel 2001: 27 (list, new com¬

bination).

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Type: M syntypes.

Habitat: Indonesia: Pulu Miang, East coast of Kaliman¬

tan. North Borneo: Landakan. “Bismarckarchipel”.

Deposited: NHMW 2230 (Stagl & Schiller, in prep.).

Castanotherium fulvicorne Pocock, 1895

Castanotherium fulvicorne Pocock, 1895: 414 (first de¬

scription)

Castanotherium fulvicorne :— Attems 1914: 150 (list)

Castanotherium fulvicorne :— Jeekel 2001: 27 (list).

Types: Holotype of unknown sex.

Habitat: Philippines: Albay.

Deposited: NHML (Online Catalogue).

Castanotherium glabratum (Newport, 1844)

Zephronia glabrata Newport, 1844: (first description)

Zephronia glabrata — Gervais 1847: 84 (list)

Zephronia glabrata :— Preudhomme de Borre 1884: 24

(list)

Zephronia glabrata :— Attems 1914: 145 (list)

Castanotherium glabrata :— Jeekel 2001: 27 (list, new

combination)

— Sphaeropoeus bicollis Karsch, 1881: 33, synonymized

with C. glabrata by Pocock 1894: 331

Sphaeropoeus bicollis :— Preudhomme de Borre 1884: 23

(list)

Sphaeropoeus bicollis :— Pocock 1894: 331 (synonymiza-

tion)

Sphaeropoeus bicollis :— Silvestri 1896a: 21 (list)

Sphaeropoeus bicollis — Attems 1914: 142 (list)

Sphaeropoeus bicollis :— Moritz & Fischer 1974: 368 (list).

Remark: Although synonymized by Pocock, the identity

of these two nominal species seems doubtful. Actual com¬

parison of the type material is necessary (Jeekel 2001).

Types: C. glabrata: holotype?, S. bicollis : M holotype

Habitat: C. glabrata: Philippines; S. bicollis : Habitat: In¬

donesia: Kalimantan, Sampit.

Deposited: C. glabrata: NHML (Online Catalogue, un¬

clear if types), S. bicollis : ZMB 714.

Castanotherium granulatum (Tomosvary, 1885)

Sphaeropoeus granulatus Tomosvary, 1885: 68 (first de¬

scription)

Sphaeropoeus granulatus — Silvestri 1896a: 21 (list)

Sphaeropoeus granulatus — Attems 1914: 144 (list)

Sphaeropoeus granulatus — Korsos 1983: 118 (list)

Castanotherium granulatum :— Attems 1936: 193 (new

combination)

Castanotherium granulatum :— Jeekel 2001: 27 (list).

©ZFMK

Thomas Wesener

Types: Syntypes.

Locality: Indonesia, Kalimantan, Matang.

Deposited: HNHM 305.60/6.

Castanotherium hirsutellum Pocock, 1895

Castanotherium hirsutellum Pocock, 1895: 414 (first de¬

scription)

Castanotherium hirsutellum.— Attems 1914: 150 (list)

Castanotherium hirsutellum :-- Wang 1951: 14 (list)

Castanotherium hirsutellum — Wang 1961: 102 (list)

Castanotherium hirsutellum — Jeekel 2001: 27 (list).

Types: Holotype of unknown sex.

Habitat: Philippines: Palawan.

Deposited: NHML (Online Catalogue).

Castanotherium hosei Pocock, 1895

Castanotherium hosei Pocock, 1895: 414 (first descrip¬

tion)

Castanotherium hosei — Attems 1914: 150 (list)

Castanotherium hosei- Hoffman 1976: 123 (description)

Castanotherium hosei— Jeekel 2001: 28 (list).

Types: Holotype of unknown sex.

Habitat: Sarawak: Baram.

Deposited: NHML (Online Catalogue).

Castanotherium humile (Silvestri, 1895)

Zephronia humilis Silvestri, 1895b: 723 (first description)

Zephronia humilis :— Attems 1914: 145 (list)

Castanotherium humile :— Jeekel 2001: 28 (list, new com¬

bination).

Types: M and F syntypes.

Habitat: Indonesia: Engano: Bua-Bua.

Deposited: IEPN.

Castanotherium ignobile (Butler, 1872)

Zephronia ignobilis Butler, 1872: 357 (first description)

Zephronia ignobilis :— Butler 1873: 180 (description)

Zephronia ignobilis — Preudhomme de Borre 1884: 24

(list)

Zephronia ignobilis :— Pocock 1894: 331 (list)

Zephronia ignobilis :— Attems 1914: 146 (list)

Castanotherium ignobile :— Jeekel 2001: 28 (list, new

combination).

Types: Holotype of unknown sex.

Habitat: Indonesia: Java.

Deposited: NHML (Online Catalogue, unclear if types).

Castanotherium indigenum (Attems, 1943)

Pulusphaera indigena Attems, 1943: 69 (first description)

Castanotherium indigenum — Jeekel 2001: 28 (list, new

combination).

Types: M holotype.

Habitat: “Borneo”.

Deposited: NHMW 2229 (Stagl & Schiller, in prep.).

Castanotherium innominatum (Butler, 1872)

Zephronia innominata Butler, 1872: 357 (first description)

Zephronia innominata :— Butler 1873: 181 (new locality)

Zephronia innominata :— Preudhomme de Borre 1884: 24

(list)

Zephronia innominata — Attems 1914: 146 (list)

Zephronia innominata — Wang 1951: 13 (list)

Zephronia innominata — Wang 1961: 101 (list)

Castanotherium innominatum — Jeekel 2001: 28 (list, new

combination).

Remark: The name of this species was already published

by Newport in 1844, but did not include a description. But¬

ler eventually validated the name in 1872.

Types: Holotype of unknown sex.

Habitat: Philippines.

Deposited: NHML (Online Catalogue).

Castanotherium insigne (Brandt, 1833)

Sphaeropoeus insignis Brandt, 1833: 200 (first descrip¬

tion)

Zephronia insignis :— Gervais 1837: 43 (list)

Sphaeropoeus insignis — Brandt 1841a: 364 (list)

Zephronia insignis — Gervais 1847: 84 (list)

Sphaeropoeus insignis — Karsch 1881: 34 (description)

Zephronia insignis :— Preudhomme de Borre 1884: 24 (list)

Sphaeropoeus insignis — Pocock 1894: 331

Castanotherium insigne — Jeekel 2001: 28 (list, new com¬

bination)

— Zephronia javanica Gervais, 1837: 43 (nomen

nudum), synonymized by Brandt 1841.

Zephronia javanica :— Guerin, 1838: 5 (locality)

Sphaerotherium javanicum :— C.L. Koch 1847: 99 (list)

Sphaerotherium javanicum :— Butler, 1873: 175 (new lo¬

cality)

Sphaerotherium javanicum :— Preudhomme de Borre

1884: 27 (list).

Remarks: Perhaps the type material of these two nominal

species, S. insignis and Z. javanica, is still in existence;

this could make it possible to finally clarify the confusion

in the classical literature. Preudhomme de Borre (1884)

viewed Z. insignis as synonym of Z ovale.

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©ZFMK

Species Catalogue Sphaerotheriida

Types: 1 M, 2 F, 2 immature syntypes.

Habitat: Indonesia: Java (Type); “East Indies”.

Deposited: ZISP.

Castanotherium laeve Carl, 1912

Castanotherium laeve Carl, 1912: 110 (first description)

Castanotherium laeve -- Attems 1914: 150 (list)

Castanotherium laeve.— Chamberlin 1920: 90 (list)

Castanotherium laeve — Jeekel 2001: 28 (list).

Types: 2 F syntypes.

Habitat: Indonesia: Matinang chain, North Sulawesi.

Deposited: MHNG.

Castanotherium lamprongse (Chamberlin, 1945)

Sphaeropoeus lamprongsis Chamberlin, 1945: 1 (first de¬

scription)

Sphaeropoeus lamprongsis :-- Wang 1967: 484 (list)

Castanotherium lamprongse — Jeekel 2001: 28 (list, new

combination).

Types: 2 M, 3 F syntypes.

Habitat: Indonesia: Sumatra, Lamprongs, Wai Lima.

Deposited: AMNH.

Castanotherium leium Chamberlin, 1921

Castanotherium leium Chamberlin, 1921: 60 (first descrip¬

tion)

Castanotherium leium :—Wang, 1951: 14 (new locality)

Castanotherium leium— hiding, 1961: 101 (new locality)

Castanotherium leium — Jeekel 2001: 29 (list).

Types: F holotype.

Habitat: Philippines: Luzon, Mt. Makiling; Mindanao,

Sungao; Zamboanga; Samar.

Deposited: MCZ 4690.

Castanotherium moderatum (Chamberlin, 1945)

Zephronia moderata Chamberlin, 1945: 1 (first descrip¬

tion)

Zephronia moderata :— Wang 1967: 484 (list)

Castanotherium moderatum :— Jeekel 2001: 29 (list, new

combination).

Types: M holotype.

Habitat: Indonesia: Java, Tjibodas.

Deposited: AMNH.

Castanotherium nigriceps (Pocock, 1894)

Zephronia nigriceps Pocock, 1894: 329 (first description)

Zephronia nigriceps — Silvestri 1895b: 723 (new locali-

ty)

Zephronia nigriceps :— Attems 1907: 105 (new locality)

Zephronia nigriceps :— Attems 1914: 146 (list)

Castanotherium nigriceps — Jeekel 2001: 29 (list, new

combination)

— Sphaerotherium kochii :— Butler 1873: 177 (misiden-

tification), NOT S. kochii Butler, 1873: 177, a synonym

of S. punctatum Brandt, synonymized by Pocock 1894:

331.

Types: M and F syntypes.

Habitat: Indonesia: Java: Bogor; Mentawei: Si-Oban. Tji¬

bodas.

Deposited: NHML (Online Catalogue, unclear if types).

Castanotherium nigromaculatum Silvestri, 1897

Castanotherium nigro-maculatum Silvestri, 1897a: 15

(first description)

Castanotherium nigromaculatum :— Attems 1914: 150

(list)

Castanotherium nigromaculatum :— Jeekel 2001: 29 (list).

Types: M holotype.

Habitat: Indonesia: Kalimantan: Amuntai.

Deposited: MTD 281.

Castanotherium obscurum Chamberlin, 1921

Castanotherium obscurum Chamberlin, 1921: 60 (first de¬

scription)

Castanotherium obscurum — Wang 1967: 484 (list)

Castanotherium obscurum :— Jeekel 2001: 29 (list).

Types: M holotype, M and F paratypes.

Habitat: Sarawak: Ladong.

Deposited: Holotype MCZ 4688, paratypes MCZ 4689.

Castanotherium ornatum Carl, 1912

Castanotherium ornatum Carl, 1912: 116 (first descrip¬

tion)

Castanotherium ornatum — Attems 1914: 150 (list)

Castanotherium ornatum :— Chamberlin 1920: 90 (list)

Castanotherium ornatum :— Jeekel 2001: 29 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sulawesi, Bontorio, South Sulawesi.

Deposited: MHNG.

Castanotherium pellitum Attems, 1932

Castanotherium pellitum Attems, 1932: 5 (first descrip¬

tion)

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Thomas Wesener

Zephronia pellita — Wang 1967: 484 (list)

Castanotheriumpellitum :— Jeekel 2001: 29 (list).

Type: M holotype.

Habitat: Indonesia: Kalimantan, Tandjoeng. Karong.

Deposited: NHMW 2226 (Stagl & Schiller, in prep.).

Castanotherium pilosum Carl, 1912

Castanotherium pilosum Carl, 1912: 114 (first description)

Castanotherium pilosum.— Attems 1914: 150 (list)

Castanotherium pilosum :- Chamberlin 1920: 90 (list)

Castanotherium pilosum :— Jeekel 2001: 29 (list).

Types: 2 F syntypes.

Habitat: Indonesia: Sulawesi, Bontorio, South Sulawesi;

Bowonglangi, South Sulawesi.

Deposited: MHNG.

Castanotherium plasoni (Attems, 1943)

Pulusphaera plasoni Attems, 1943: 70 (first description)

Castanotherium plasoni — Jeekel 2001: 29 (list, new com¬

bination).

Types: M holotype.

Habitat: Indonesia: Java, Samarang.

Deposited: NHMW 2232 (Stagl & Schiller, in prep.).

Castanotherium pyrrhomelanum (Attems, 1897)

Zephroniapyrrhomelana Attems, 1897: 480 (first descrip¬

tion)

Zephronia pyrrhomelana :— Attems 1914: 146 (list)

Castanotherium pyrrhomelanum :— Jeekel 2001: 29 (list,

new combination).

Types: Syntypes of unknown sex.

Habitat: Sarawak: Baram river.

Deposited: NHMW 2234 (Stagl & Schiller, in prep.).

Castanotherium rufipes (Pocock, 1894)

Zephronia rufipes Pocock, 1894: 331 (first description)

Zephronia rufipes (“ruficeps”):—Attems 1898: 509 (new

locality)

Zephronia rufipes (“ruficeps”):- Attems 1903: 64 (new

locality)

Zephronia rufipes (“ruficeps”):— Attems 1914: 147 (list)

Castanotherium rufipes — Jeekel 2001: 29 (list, new com¬

bination).

Type: M holotype.

Habitat: Indonesia: Java, Tjibodas.

Deposited: NHML (online catalogue).

Castanotherium simplex (Carl, 1909)

Sphaeropoeus simplex Carl, 1909: 250 (first description)

Castanotherium simplex :— Attems 1914: 150 (list)

Castanotherium simplex :— Jeekel 2001: 29 (list).

Types: M and F syntypes.

Habitat: Indonesia: Java: Prigen.

Deposited: MHNG.

Castanotherium sparsepunctatum Carl, 1912

Castanotherium sparsepunctatum Carl, 1912: 120 (first

description)

Castanotherium sparsepunctatum :— Attems 1914: 150

(list)

Castanotherium sparsepunctatum :— Chamberlin 1920: 91

(list)

Castanotherium sparsepunctatum :— Jeekel 2001: 30

(list).

Types: F holotype.

Habitat: Indonesia: Sulawesi, Bowonglangi, South Su¬

lawesi.

Deposited: MHNG.

Castanotherium stellatum Carl, 1912

Castanotherium stellatum Carl, 1912: 122 (first descrip.)

Castanotherium stellatum — Attems 1914: 150 (list)

Castanotherium stellatum :— Chamberlin 1920: 91 (list)

Castanotherium stellatum — Jeekel 2001: 30 (list).

Types: F syntypes.

Habitat: Indonesia: Sulawesi, Loka, South Sulawesi.

Deposited: MHNG.

Castanotherium sulcicollis (Karsch, 1881)

Sphaeropoeus sulcicollis Karsch, 1881: 32 (first descrip¬

tion)

Zephronia sulcicollis :— Preudhomme de Borre 1884: 25

(list)

Sphaeropoeus sulcicollis :— Tomosvary 1885: 68 (new lo¬

cality)

Sphaeropoeus sulcicollis :— Silvestri 1896a: 21 (list)

Sphaeropoeus sulcicollis :— Attems 1914: 143 (list)

Sphaeropoeus sulcicollis :—Wang 1951: 13 (list)

Sphaeropoeus sulcicollis :— Wang 1961: 101 (list)

Sphaeropoeus sulcicollis :— Moritz & Fischer 1974: 370

(list)

Castanotherium sulcicollis — Jeekel 2001: 30 (list, new

combination).

Remarks: The type series contains at least three different

species (Moritz & Fischer 1974: 370).

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©ZFMK

Species Catalogue Sphaerotheriida

Types: M and F syntypes.

Habitat: Indonesia: Java: Palabuan; Djakarta; Kaliman¬

tan, Singkawang; Benkajang;

Montrado. Philippines: Luzon.

Deposited: ZMB 195, ZMB 495, ZMB 912, ZMB 913,

ZMB 587, ZMB 588, ZMB 589, ZMB 591, ZMB 597.

Cast an oth erium suspectum C arl, 1912

Castanotherium suspectum Carl, 1912: 109 (first descrip¬

tion)

Castanotherium suspectum.— Attems 1914: 150 (list)

Castanotherium suspectum :— Chamberlin 1920: 90 (list)

Castanotherium suspectum :— Jeekel 2001: 30 (list).

Remarks: According to Carl (1912: 109) potentially the

unknown female of C. distinctum.

Types: F holotype (damaged).

Habitat: Indonesia: Sulawesi, Mpane, Gulf of Tontini,

Central Sulawesi.

Deposited: MHNG.

Castanotherium tivium (Chamberlin, 1945)

Zephronia tivia Chamberlin, 1945: 2 (first description)

Castanotherium tivium — Jeekel 2001: 30 (list, new com¬

bination).

Types: M holotype.

Habitat: Indonesia: Sumatra, Lamprongs, Wai Lima.

Deposited: AMNH.

Castanotherium tricollis (Karsch, 1881)

Sphaeropoeus tricollis Karsch, 1881: 32 (first description)

Zephronia tricollis :— Preudhomme de Borre 1884: 25 (list)

Sphaeropoeus tricollis :— Tomosvary 1885: 68 (new local¬

ity)

Sphaeropoeus tricollis :— Attems 1914: 144 (list)

Sphaeropoes [sic!] tricollis — Wang 1967: 484 (list)

Sphaeropoeus tricollis :— Moritz & Fischer 1974: 370 (list)

Castanotherium tricollis :— Jeekel 2001: 30 (list, new com¬

bination).

Types: 16 M, 7 F syntypes (Moritz & Fischer 1974: 370).

Habitat: Indonesia: Sumatra: Lahat; Tibingtingi; Java.

Deposited: ZMB 574, ZMB 914, ZMB 2191.

Castanotherium variegatum (Attems, 1943)

Pulusphaeravariegata Attems, 1943: 68 (first description)

Pulusphaera variegata :— Wang 1967: 485 (list)

Castanotherium variegatum :— Jeekel 2001: 30 (list, new

combination).

Types: M holotype.

Habitat: Sarawak: Baram distr.

Deposited: NHMW 2228 (Stagl & Schiller, in prep.).

Castanotherium volzi (Carl, 1906)

Sphaeropoeus volzi Carl, 1906: 233 (first description)

Castanotherium volzi :— Attems 1914: 151 (list).

Castanotherium volzi — Jeekel 2001: 30 (list).

Types: M and F syntypes.

Habitat: Indonesia: Sumatra, Palembang; Indragiri, Lahat.

Deposited: MHNG.

Castanotherium whiteheadi Pocock, 1895

Castanotherium whiteheadi Pocock, 1895: 414 (first de¬

scription)

Castanotherium whiteheadi— Attems 1914: 151 (list)

Castanotherium whiteheadi — Jeekel 2001: 30 (list).

Type: Holotype of unknown sex.

Habitat: North Borneo: Kina Balu.

Deposited: NHML (online catalogue).

Castanotheroides Chamberlin. 1921

Castanotheroides Chamberlin, 1921: 61 (first description);

Wang 1951: 14 (list); Wang 1961: 102 (list); Jeekel 1971:

24 (list); Jeekel 1974: 46 (classification); Hoffman 1976:

120 (list); Hoffman 1980: 64 (list); Jeekel 2001: 30 (list,

re-definition).

Synonyms:

— Luzonosphaera Wang, 1951: 14, synonymized with

Castanotheroides by Jeekel 2001: 30. Wang 1961: 103;

Jeekel 1971: 26; Jeekel 1974: 46; Hoffman 1976: 120;

Hoffman 1980: 64; Mauries 2001: 141.

Tvpe-species : Luzonosphaera philippina Wang, 1951.

Description: Anterior and posterior telopods apparently

similar to Castanotherium , but antennomere 6 cylindrical

and in both sexes with only four apical cones.

Type-species: Castanotherium porosum Pocock.

Species included (3)

Castanotheroidesphilippinus (Wang, 1951) (Philippines)

Castanotheroides porosus (Pocock, 1895) (Philippines)

Castanotheroides sinuatus (Butler, 1872) (Borneo)

Castanotheroides philippinus (Wang, 1951)

Luzonosphaera philippina Wang, 1951: 15 (first descrip.)

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Thomas Wesener

Luzonosphaeraphilippina — Wang 1961: 103 (descrip.)

Castanotheroidesphilippinus — Jeekel 2001: 31 (list, new

combination).

Types: M holotype, 1 M and 1 F paratypes.

Habitat: Philippines: Luzon: Mt. Makiling.

Deposited: Unknown.

Castanotheroides porosus (Pocock, 1895)

Castanotherium porosnm Pocock, 1895: 414 (first descrip¬

tion)

Castanotherium porosum — Attems 1914: 150 (list)

Castanotheroides porosus :— Chamberlin 1921: 61 (list)

Castanotheroides porosus.— Wang 1951: 14 (list)

Castanotheroides porosus.— Wang 1961: 102 (list)

Castanotheroides porosus :— Jeekel 2001: 31 (list).

Types: Syntypes of unknown sex.

Habitat: Philippines.

Deposited: NHML (Online Catalogue), unclear of types.

Castanotheroides sinuatus (Butler, 1872)

Sphaerotherium sinuatum Butler, 1872: 359 (first descrip.)

Sphaerotherium sinuatum :— Butler 1873: 174 (list)

Sphaerotherium sinuatum — Preudhomme de Borre 1884:

28 (list)

Sphaerotherium sinuatum — Attems 1914: 141 (list)

Castanotheroides sinuatus :— Jeekel 2001: 31 (list, new

combination).

Types: Holotype of unknown sex.

Habitat: Sarawak.

Deposited: NHML.

Sechelliosoma Mauries. 1980

Sechelliosoma Mauries, 1980: 164 (subgenus of Cylio-

somella ); Golovatch & Korsos 1992: 2 (new localities);

Shelley et al. 2000: 13 (list); Mauries 2001: 141 (list); We¬

sener & VandenSpiegel 2009: 548 (morphological phylo¬

genetic analysis).

Description: Anterior and posterior telopods with three

podomeres distal to syncoxite, both pairs strongly elon¬

gated and slender. Antennomere 6 cylindrical, disc with

only four apical cones. Mandible inner tooth only consist¬

ing of two combs. <20 mm long.

Type-species: Sphaerotherium forcipatum Brolemann,

1896.

Monotypic

Sechelliosoma forcipatum (Brolemann, 1896)

Sphaerotherium forcipatum Brolemann, 1896: 529 (first

description)

Cyliosomella {Sechelliosoma)furcipatum [sic!]: — Mauries

1980: 164 (new subgenus, new localities)

Sechelliosoma forcipatum :— Golovatch & Korsos 1992:

2 (raised to genus)

Sechelliosoma forcipatum — Wesener & VandenSpiegel

2009: 548 (phylogenetic analysis)

Sechelliosoma forcipatum :— Golovatch & Gerlach 2010:

387 (key).

Types: 1 M, 6 F syntypes.

Habitat: Seychelles: Mahe, Silhouette, Felicite, Marianne.

Deposited: MNHN.

Subfamily Rajasphaerinae Hoffman

Hoffman, 1976: 120; Hoffman 1980: 64; Jeekel 2001: 31;

Mauries 2001: 141.

Remark: A subfamily Rajasphaerinae was mentioned by

Verhoeff (1937) but was never diagnosed (Jeekel 2001).

Genus Rajasphaera Attems. 1935

Rajasphaera Attems, 1935: 126 (first description); Attems

1943: 62 (list); Jeekel 1971: 28 (list); Jeekel 1974: 46

(classification); Hoffman 1976: 12 (redescription); Hoff¬

man 1980: 64 (list); Jeekel 2001: 31 (list); Mauries 2001:

141 (list).

Description: Anterior telopod apparently with three

podomeres distal to syncoxite. Posterior telopods unique,

telopoditomere 1 massively enlarged and swollen, while

telopoditomere 2 is also quite large, with the immovable

finger almost fused to its basal parts. Telopoditomere 3

very slender. Male gonopore located almost mesally in¬

stead of posteriorly. Antennomere 6 strongly axe-shaped.

Up to 50 mm long.

Type-species: Rajasphaera montana Attems.

Monotypic

Rajasphaera montana Attems, 1935

Rajasphaera montana Attems, 1935: 132 (first descrip¬

tion)

Rajasphaera montana:— Hoffman 1976: 121 (redescrip¬

tion)

Rajasphaera montana :— Jeekel 2001: 31 (list)

NOT Sphaeropoeus montanus Karsch, 1881: 31, now

Zephronia montana.

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Species Catalogue Sphaerotheriida

Types: M holotype (telopods missing).

Habitat: Indonesia: Kalimantan, Mt. Raja.

Deposited: ZMB.

Nomina dubia of the Zephroniidae

Sphaerotherium nebulosum Butler, 1875

Sphaerotherium nebulosum Butler, 1875: 165 (first de¬

scription)

Sphaerotherium nebulosum .— Preudhomme de Borre

1884: 27 (list)

Types: 1 holotype of unknown sex.

Locality: Mongolia (?)

Deposited: NHML (presumably lost).

Family Sphaerotheriidae Brandt. 1833

Sphaerotheriidae Brolemann 1913: 79 (first separation of

the Sphaerotheriida into Sphaerotheriidae and

‘Sphaeropoeidae’); Attems 1914: 140 (list); Jeelcel 1974:

45 (classification); VandenSpiegel et al. 2002: 143 (revi¬

sion); Wesener & VandenSpiegel, 2009: 561 (morpholog¬

ical phylogenetic analysis); Wesener et al. 2010b: 1186

(molecular phylogenetic analysis); Wesener et al. 2011:

967 (stridulation)

Remarks: Before 2009 (Wesener & VandenSpiegel) the

family Sphaerotheriidae was a paraphyletic group contain¬

ing the genera Procyliosomatidae, Cyliosomatidae and

Arthrosphaeridae in addition to the current species.

Genera included (2)

Sphaerotherium Brandt, 1833 (eastern South Africa, Mo-

cambique, Zimbabwe)

Kylindotherium Attems, 1926 (South Africa)

Type genus: Sphaerotherium Brandt, 1833

55 species + 1 nomina dubia.

Genus Sphaerotherium Brandt 1833

(Figs 12A-C)

Sphaerotherium Brandt, 1833: 198 (first description);

Brandt 1841a: 360 (new species); Brandt 1841b: 177 (new

species); Koch 1847: 99 (new species); Deshayes & Milne

Edwards 1838: 47 (list); Lucas 1842: 522 (new species);

Newport 1844: 10 (list); Koch 1863: 40 (description); Po-

rat 1872: 5 (new species); Butler 1873:175 (new species);

Karsch 1881: 20 (new species); Bourne 1885: 162 (de¬

scription); Pocock 1895 (redefinition); Attems 1908: 423

(new species); Silvestri 1910: 191 (new species); Attems

1914: 140 (list); Attems 1926 (key); Attems 1928: 220

(new species); Attems 1934 (new species); Warren 1934:

382 (description); Lawrence 1953: 77 (list); Manton 1954:

299 (morphology); Schubart 1958: 48 (new species);

Lawrence 1966: 242 (new species); Haacker 1969: 16 (be¬

haviour); Haacker & Fuchs 1972: 191 (behaviour);

Haacker 1974: 317 (behaviour); Jeekel 1974: 45 (classi¬

fication); Raseroka 1975: 52 (predation); Hoffman 1980

(list); Jocque 1984: 67 (new species); Lawrence 1987: 113

(list); Alderweireldt 1997: 53 (new species); Hamer 1998:

15 (list); Jones & jones 2000:234 (medicine); Herbert

2000 (predation by slugs); Golovatch & Hoffman 2001:

238 (list); VandenSpiegel et al. 2002: 145 (revision); Her¬

bert et al. 2003: 4 (medicine); Redi et al. 2005: 287 (ecol¬

ogy); Wesener & VandenSpiegel, 2009: 561 (morpholog¬

ical phylogenetic analysis); Wesener et al. 2010b: 1186

(molecular phylogenetic analysis); Wesener et al. 2011:

967 (stridulation).

Synonyms:

— Bournellum de Saussure & Zehntner, 1902: 20, for¬

mally synonymized under Sphaerotherium by Vanden¬

Spiegel et al. 2002: 145. Jeekel 1971: 23 (list).

Type species : Sphaerotherium retusum Koch, 1847: 99,

subsequently designated by Jeekel 1971: 23, syn¬

onymized under Sphaerotherium dorsale (Gervais,

1847) by Attems 1928: 227.

— Eubournellum Attems, 1908: 426, synonymized by

Jeekel 1971: 25, as junior synonym of Bournellum de

Saussure & Zehntner, 1902.

Type species : Sphaerotherium retusum Koch, 1847: 99,

subsequently designated by Jeekel 1971: 23, syn¬

onymized under Sphaerotherium dorsale (Gervais,

1847) by Attems 1928: 227.

— Natalobelum Verhoeff, 1924: 59 synonymized with

Sphaerotherium by VandenSpiegel et al. 2002: 145.

Jeekel 1971: 27 (list).

Type species : Sphaerotheriumpunctulatum Brandt, 1841,

by monotypy.

— Neobournellum Attems, 1908: 426, synonymized with

Sphaerotherium by VandenSpiegel et al. 2002: 145.

Type species : Bournellum trichopygum (Attems, 1908:

424), by monotypy, synonymized under Sphaerotheri¬

um compressum Brandt, 1833: 198 by VandenSpiegel

et al. 2002: 151.

— Oligaspis Wood, 1865: 173 (first description), syn¬

onymized with Sphaerotherium by VandenSpiegel et al.

2002: 145. Jeekel 1971: 27 (list).

Type species : Oligaspispunctipes Wood, 1865, by mono¬

typy, synonymized under Sphaerotherium rotundatum

Brandt, 1833: 198 by VandenSpiegel et al. 2002: 168.

— Tetraconosoma Verhoeffj 1924: 57, synonymized with

Sphaerotherium by VandenSpiegel et al. 2002: 145.

Jeekel 1971: 29 (list).

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Type species : Sphaerotherium weberii Silvestri, 1910, by

subsequent designation by Jeekel 1971: 29.

Description: See family key above.

Type species: Sphaerotherium rotundatum Brandt, 1833

by subsequent designation of Pocock, 1895: 410.

Included species (54)

Sphaerotherium alticola Schubart, 1958

Sphaerotherium apicale Silvestri, 1910

Sphaerotherium boerium Silvestri, 1910

Sphaerotherium capense Schubart, 1958

Sphaerotherium cinctellum Silvestri, 1910

Sphaerotherium civicum Attems, 1928

Sphaerotherium commune Attems, 1928

Sphaerotherium compressum Brandt, 1833

Sphaerotherium coniferum Silvestri, 1910

Sphaerotherium convexitarsum Silvestri, 1910

Sphaerotherium dicrothrix Attems, 1928

Sphaerotherium dinogonum Silvestri, 1910

Sphaerotherium dorsale (Gervais, 1847)

Sphaerotherium dorsaloide Silvestri, 1910

Sphaerotherium elongatum Brandt, 1833

Sphaerotherium eremita Attems, 1928

Sphaerotherium eucalyptophilum Schubart, 1958

Sphaerotherium fulvum Schubart, 1958

Sphaerotherium giganteum Porat, 1872

Sphaerotherium granulatum Pocock, 1895

Sphaerotherium hanstroemi Schubart, 1958

Sphaerotherium intermedium Porat, 1872

Sphaerotherium kitharistes Attems, 1928

Sphaerotherium krugeri Lawrence, 1966

Sphaerotherium lichtensteinii Brandt, 1833

Sphaerotherium mahaium Schubart, 1958

Sphaerotherium millepunctatum Attems, 1928

Sphaerotherium modestum Attems, 1928

Sphaerotherium narcissei Alderweireldt, 1997

Sphaerotherium nigritarse Schubart, 1958

Sphaerotherium obtusum C. L. Koch, 1847

Sphaerotherium perbrincki Schubart, 1958

Sphaerotherium permodestum Silvestri, 1910

Sphaerotherium pinnatum Schubart, 1958

Sphaerotherium plagiarium Silvestri, 1910

Sphaerotherium punctatum Brandt, 1833

Sphaerotherium punctulatum Brandt, 1841

Sphaerotherium rotundatum Brandt, 1833

Sphaerotherium rudebecki Schubart, 1958

Sphaerotherium rugulosum Brandt, 1841

Sphaerotherium selindum Alderweireldt, 1997

Sphaerotherium similare Schubart, 1958

Sphaerotherium solitarium Attems, 1928

Sphaerotherium spinatum Silvestri, 1910

Sphaerotherium steppense Schubart, 1958

Sphaerotherium subdorsale Silvestri, 1910

Sphaerotherium submite Silvestri, 1910

Sphaerotherium tenuitarse Silvestri, 1910

Sphaerotherium tomentosum Schubart, 1958

Sphaerotherium transzambeziacum Jocque, 1984

Sphaerotherium tuberosum Attems, 1928

Sphaerotherium tzitzikamum Schubart, 1958

Sphaerotherium weberii Silvestri, 1910

Sphaerotherium zuluense Schubart, 1958

Sphaerotherium alticola Schubart, 1958

Sphaerotherium (Tetraconosoma) alticola Schubart, 1958:

69 (first description)

Sphaerotherium (Tetraconosoma) alticola.— Hamer 1998:

15 (list)

Sphaerotherium alticola :-- VandenSpiegel et al. 2002: 174

(list).

Types: F holotype.

Habitat: Lesotho: Mt. Morosi, 15 miles NE Quthing, top

plateau.

Deposited: MZLU.

Sphaerotherium apicale Silvestri, 1910

Sphaerotherium apicale Silvestri, 1910: 217 (first descrip¬

tion)

Sphaerotherium apicale — Attems 1928: 229 (list)

Sphaerotherium (Sphaerotherium) apicale :-- Schubart

1958: 93 (list)

Sphaerotherium apicale — Lawrence 1966: 242 (list)

Sphaerotherium (Sphaerotherium) apicale — Hamer 1998:

15 (list)

Sphaerotherium apicale — VandenSpiegel et al. 2002: 174

(list).

Types: M holotype.

Habitat: Mozambique: Maputo.

Deposited: IEPN or ZMUH.

Sphaerotherium boerium Silvestri, 1910

Sphaerotherium boerium Silvestri, 1910: 219 (first de¬

scription)

Sphaerotherium boerium :— Attems 1928: 229 (list)

Sphaerotherium (Sphaerotherium) boerium :— Schubart

1958: 93 (list)

Sphaerotherium (Sphaerotherium) boerium — Hamer

1998: 15 (list)

Sphaerotherium boerium — VandenSpiegel et al. 2002:

174 (list).

Types: M holotype.

Habitat: South Africa: Gauteng: Pretoria.

Deposited: IEPN.

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Species Catalogue Sphaerotheriida

Sphaerotherium capense Schubart, 1958

Sphaerotherium (Tetraconosoma) capense Schubart,

1958: 48 (first description)

Sphaerotherium (Tetraconosoma) capense.— Hamer 1998:

15 (list)

Sphaerotherium capense :— VandenSpiegel et al. 2002: 174

(list)

Sphaerotherium capense :— Pryke & Samways 2010: 251

(ecology)

— Sphaerotherium monticola Pocock, 1895: 410 (first

description), synonymy under S. capense proposed in

Schubart 1958: 50

Sphaerotherium monticola :— Attems 1928: 236 (list)

Sphaerotherium monticola:— Schubart 1958: 101 (list)

Sphaerotherium monticola:— VandenSpiegel et al. 2002:

180 (list).

Types: M holotype, 1 F paratype.

Habitat: South Africa: W. Cape: Table Mtn, Blinkwater

Ravine.

Deposited: MZLU.

Sphaerotherium cinctellum Silvestri, 1910

Sphaerotherium cinctellum Silvestri, 1910: 191 (first de¬

scription)

Sphaerotherium cinctellum - Attems 1928: 222 (descrip¬

tion)

Sphaerotherium cinctellum :— Attems 1937: 464 (descrip¬

tion)

Sphaerotherium (Tetraconosoma) cinctellum — Schubart

1958: 86 (description)

Sphaerotherium cinctellum :— Haacker & Fuchs 1972: 191

(behaviour)

Sphaerotherium cinctellum :— Lawrence 1987: 113 (list)

Sphaerotherium (Tetraconosoma) cinctellum — Hamer

1998: 16 (list)

Sphaerotherium cinctellum :— VandenSpiegel et al. 2002:

174 (list)

Sphaerotherium cinctellum :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Types: F holotype.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium civicum Attems, 1928

Sphaerotherium civicum Attems, 1928: 231 (first descrip.)

Sphaerotherium (Sphaerotherium) civicum — Schubart

1958: 88 (list);

Sphaerotherium (Sphaerotherium) civicum — Hamer

1998: 16 (list)

Sphaerotherium civicum :— VandenSpiegel et al. 2002: 174

(list).

Types: M and F syntypes.

Habitat: South Africa: E. Cape: Burgersdorp.

Deposited: SAMC, NHMW 3953 (Stagl & Schiller, in

prep.).

Sphaerotherium commune Attems, 1928

Sphaerotherium commune Attems, 1928: 218 (first de¬

scription)

Sphaerotherium (Tetraconosoma) commune — Schubart

1958: 58 (list)

Sphaerotherium (Tetraconosoma) commune — Hamer

1998: 16 (list)

Sphaerotherium commune :— VandenSpiegel et al. 2002:

174 (list).

Types: Numerous M and F syntypes.

Habitat: South Africa, numerous localities, no type local¬

ity specified.

Deposited: SAMC; NHMW 2199 (Stagl & Schiller, in

prep.).

Sphaerotherium compressum Brandt, 1833

Sphaerotherium compressum Brandt. 1833: 198 (first de¬

scription)

Zephronia compressa :— Gervais: 1837: 9 (list)

Zephronia compressa :— Deshayes & Milne Edwards 1838:

47 (list)

Sphaerotherium compressum — Brandt 1841a: 360 (de¬

scription)

Sphaerotherium compressum :— Brandt 1841b: 176 (de¬

scription)

Zephronia compressa :— Lucas 1842: 521 (description)

Zephronia compressa :— Gervais 1847: 77 (description)

Sphaerotherium compressum :— Koch 1847: 99 (list)

Sphaerotherium compressum :— Koch 1863: 45 (descrip.)

Sphaerotherium compressum — Butler 1873: 175 (list)

Sphaerotherium compressum :— Preudhomme de Borre

1884: 26 (list)

Sphaerotherium compressum :— Silvestri 1910: 199 (de¬

scription)

Sphaerotherium compressum :— Moritz & Fischer 1973:

368 (list)

Sphaerotherium compression:— Golovatch & Hoffman

2001: 238 (list)

Sphaerotherium compressum :— VandenSpiegel et al.

2002: 151 (redescription)

Sphaerotherium compression:— Wesener & Vanden¬

Spiegel 2009: 548 (morpholog. phylogenetic analysis)

— Sphaerotherium Klugii [sic!] Brandt, 1841a: 361 (first

description), synonymized with S. compressum by Van¬

denSpiegel et al. 2002: 151

Sphaerotherium Klugii [sic!]:— Brandt 1841 b: 177 (des¬

cription)

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Zephronia Klugii [sic!]:— Gervais 1847: 79 (description)

Sphaerotherium klugii.— Butler 1873: 176 (description)

Sphaerotherium Klugii [sic!]:— Preudhomme de Borre

1884: 27 (list)

Sphaerotherium compressum klugii— Silvestri 1910: 201

(description)

Sphaerotherium klugii— Golovatch & Hoffman 2001: 238

(list)

— Sphaerotherium repulsum Karsch, 1881: 28 (first des¬

cription), synonymized with S. compressum by Vanden-

Spiegel etal. 2002: 151

Sphaerotherium repulsum — Moritz & Fischer 1973: 369

(list)

— Bournellum trichopygum Attems, 1908: 424 (first de¬

scription), synonymized with S. compressum by Vanden-

Spiegel etal. 2002: 151

Sphaerotherium trichopygum — Attems 1928: 221 (de¬

scription)

Sphaerotherium trichopygum :— Moritz & Fischer 1973:

370 (list)

Sphaerotherium (Tetraconosoma) trichopygum—

Schubart, 1958: 56 (description)

Sphaerotherium (Tetraconosoma) trichopygum :— Hamer

1998: 19 (list)

— Sphaerotherium ancillare Attems, 1928: 225 (first de¬

scription), synonymized with S. compressum by Vanden-

Spiegel et al. 2002: 151

Sphaerotherium (Tetraconosoma) ancillare :— Schubart

1958: 86 (list)

Sphaerotherium (Tetraconosoma) ancillare — Hamer

1998: 15 (list).

Types: M lectotype (designated by VandenSpiegel et al.

2002: 151), 2 F paratypes.

Habitat: South Africa: ‘Promontorium Bonae Spei’

(Cape).

Deposited: Lectotype and 1 paratype, ZMB 62; 1 paraty-

pe ZISP

Sphaerotherium coniferum Silvestri, 1910

Sphaerotherium coniferum Silvestri, 1910: 213 (first de¬

scription)

Sphaerotherium coniferum :— Attems 1928: 230 (descrip¬

tion)

Sphaerotherium (Sphaerotherium) coniferum :— Schubart

1958: 93 (list)

Sphaerotherium (Sphaerotherium) coniferum :— Hamer

1998: 16 (list)

Sphaerotherium coniferum :— VandenSpiegel et al. 2002:

175 (list).

Types: F holotype.

Habitat: Mozambique: Maputo.

Deposited: IEPN or ZMUH.

Sphaerotherium convexitarsum Silvestri, 1910

Sphaerotherium convexitarsum Silvestri, 1910: 198 (first

description)

Sphaerotherium convexitarsum :— Attems 1928: 217 (list)

Sphaerotherium (Tetraconosoma) convexitarsum—

Schubart 1958: 50 (list)

Sphaerotherium (Tetraconosoma) convexitarsum :— Hamer

1998: 16 (list)

Sphaerotherium convexitarsum :— VandenSpiegel et al.

2002: 175 (list)

Sphaerotherium convexitarsum — Wesener et al. 2011: 967

(stridulation behaviour).

Types: M and F syntypes.

Habitat: South Africa: E. Cape: Port Elizabeth.

Deposited: IEPN.

Sphaerotherium dicrothrix Attems, 1928

Sphaerotherium dicrothrix Attems, 1928: 232 (first de¬

scription)

Sphaerotherium (Sphaerotherium) dicothrix [sic!]:—

Schubart 1958: 87 (list)

Sphaerotherium (Sphaerotherium) dicrothrix:- Hamer

1998: 16 (list)

Sphaerotherium dicrothrix:- VandenSpiegel et al. 2002:

175 (list).

Types: M and F syntypes.

Habitat: South Africa: LP: Acomhoek.

Deposited: SAMC, NHMW 2200 (Stagl & Schiller, in

prep.).

Sphaerotherium dinogonum Silvestri, 1910

Sphaerotherium dinogonum Silvestri, 1910: 210 (first de¬

scription)

Sphaerotherium dinogonum — Attems 1928: 226 (list)

Sphaerotherium (Sphaerotherium) dinogonum :— Schubart

1958: 94 (list)

Sphaerotherium (Sphaerotherium) dinogonum — Hamer

1998: 16 (list)

Sphaerotherium dinogonum — VandenSpiegel et al. 2002:

175 (list).

Types: M and F syntypes.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium dorsale (Gervais, 1847)

Zephronia dorsalis Gervais, 1847: 79 (first description)

Sphaerotherium dorsale :— Porat 1872: 5 (description)

Sphaerotherium dorsale :— Preudhomme de Borre 1884:

26 (list)

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Species Catalogue Sphaerotheriida

Sphaerotherium dor sale.— Silvestri 1910: 205 (descrip.)

Sphaerotherium dorsale — Attems 1928: 227 (description)

Sphaerotherium dorsale - Attems 1934: 464 (description)

Sphaerotherium dorsale - Lawrence 1953: 77 (list)

Sphaerotherium dorsale — Lawrence 1987: 110 (list)

Sphaerotherium dorsale :— Manton 1954: 299 (description)

Sphaerotherium dorsale :— Schubart 1958: 98 (description)

Sphaerotherium dorsale — Haacker 1968: 454 (behaviour)

Sphaerotherium dorsale :— Haacker & Fuchs 1972: 192

(behaviour)

Sphaerotherium dorsale :— Hamer 1998: 16 (list)

Sphaerotherium dorsale — VandenSpiegel et al. 2002: 154

(redescription)

Sphaerotherium dorsale — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaerotherium dorsale — Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium dorsale :—Armstrong & Hamer 2015: 153

(list)

— Zephroniapuverea White, 1859: 405 (first descrip.),

synonymized with S. dorsale by Schubart 1958: 98.

— Sphaerotherium retusum Koch, 1847: 99 (first descrip¬

tion), synonymized with S. dorsale by Attems 1928: 227

Sphaerotherium retusum :— Koch 1863: 42 (description)

Sphaerotherium retusum :— Bourne 1885: 162 (descrip.)

Bournellum retusum :— De Saussure & Zehntner 1902: 22

(description).

Types: F holotype.

Habitat: Unknown.

Deposited: MNHN.

Sphaerotherium dorsaloide Silvestri, 1910

Sphaerotherium dorsaloide Silvestri, 1910: 189 (first de¬

scription)

Sphaerotherium dorsaloide :— Attems 1928: 225 (descrip.)

Sphaerotherium dorsaloide :— Lawrence 1953: 88 (de¬

scription)

Sphaerotherium (Tetraconosoma) dorsaloide :— Schubart

1958: 80 (description)

Sphaerotherium (Tetraconosoma) dorsaloide :— Hamer

1998: 16 (list)

Sphaerotherium dorsaloide :— VandenSpiegel et al. 2002:

175 (list)

Sphaerotherium dorsaloide :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaerotherium dorsaloide — Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium dorsaloides [sic!]:— Armstrong & Hamer

2015: 153 (list).

Types: M and F syntypes.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium elongatum Brandt, 1833

Sphaerotherium elongatum Brandt, 1833: 199 (first de¬

scription)

Zephronia elongata — Gervais 1837: 9 (list)

Zephronia elongata :— Gervais 1847: 80 (description)

Zephronia elongata :— Deshayes & Milne Edwards 1838:

47 (list)

Sphaerotherium elongatum :— Brandt 1841a: 362 (descrip¬

tion)

Sphaerotherium elongatum — Brandt 1841b: 178 (descrip¬

tion)

Zephronia elongata — Lucas 1842: 522 (description)

Sphaerotherium elongatum — Newport 1844: 10 (list)

Sphaerotherium elongatum :— Koch 1847: 98 (list)

Sphaerotherium elongatum :— Koch 1863: 32 (description)

Sphaerotherium elongatum :— Butler 1873: 176 (list)

Sphaerotherium elongatum :— Preudhomme de Borre

1884: 26 (list)

Sphaerotherium elongatum — Moritz & Fischer 1973: 368

(list)

Sphaerotherium elongatum :— Golovatch & Hoffman

2001: 238 (list)

Sphaerotherium elongatum — VandenSpiegel et al. 2002:

156 (redescription)

— Sphaerotherium microstictum Brandt, 1841a: 362 (first

description), synonymized under 5. elongatum by Van¬

denSpiegel etal. 2002: 156

Sphaerotherium microstictum — Brandt 1841b: 178 (de¬

scription)

Zephronia microsticta — Gervais 1847: 80 (description)

Sphaerotherium microstictum — Butler 1873: 173 (descrip¬

tion)

Sphaerotherium microstictum — Preudhomme de Borre

1884: 27 (list)

Sphaerotherium microstictum :— Golovatch & Hoffman

2001: 239 (list)

Sphaerotherium microstictum — VandenSpiegel et al.

2002: 156 (synonymization).

— Sphaerotherium insulanum Karsch, 1881: 30 (first de¬

scription), synonymized under S. elongatum by Wesen¬

er &Wagele 2008: 79

Sphaerotherium insulanum — Preudhomme de Borre

1884: 26 (list)

Sphaerotherium insulanum :— Jeekel 1999: 14 (list).

Types: M lectotype, 4 F paralectotype (designated by Van¬

denSpiegel et al. 2002: 156).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: Lectotype and 2 paralaectotypes ZMB 60, 1

paralectotype ZMB 59, 1 paralectotype ZISP

Sphaerotherium eremita Attems, 1928

Sphaerotherium eremita Attems, 1928: 229 (first descrip¬

tion)

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Thomas Wesener

Sphaerotherium (Sphaerotherium) eremita:— Schubart

1958: 93 (list)

Sphaerotherium (Sphaerotherium) eremita.— Hamer 1998:

17 (list)

Sphaerotherium eremita :— VandenSpiegel et al. 2002: 176

(list).

Types: F holotype.

Habitat: South Africa: W. Cape: Table Mtn, Newlands.

Deposited: SAMC; NHMW 72/229 (Stagl & Schiller, in

prep.).

Sphaerotherium eucalyptophilum Schubart, 1958

Sphaerotherium (Tetraconosoma) eucalyptophilum

Schubart, 1958: 60 (first description)

Sphaerotherium (Tetraconosoma) eucalyptophilum :—

Hamer 1998: 17 (list)

Sphaerotherium eucalyptophilum — VandenSpiegel et al.

2002: 176 (list).

Types: M holotype, 1 F paratype.

Habitat: South Africa: LP: Soutpansberg, Punch Bowl 1.6

miles NNE Louis Trichardt.

Deposited: MZLU.

Sphaerotherium fulvum Schubart, 1958

Sphaerotherium (Tetraconosoma) fulvum Schubart, 1958:

67 (first description)

Sphaerotherium (Tetraconosoma) fulvum — Hamer 1998:

17 (list)

Sphaerotherium fulvum :— VandenSpiegel et al. 2002: 176

(list).

Types: M holotype.

Habitat: South Africa: E. Cape: Zuurberg Pass, 15 miles

N Addo.

Deposited: MZLU.

Sphaerotherium giganteum Porat, 1872

(Fig. 12 A)

Sphaerotherium giganteum Porat, 1872: 8 (first descrip.)

Sphaerotherium giganteum — Preudhomme de Borre

1884: 26 (list)

Sphaerotherium giganteum — Silvestri 1910: 212 (descrip¬

tion)

Sphaerotherium giganteum :— Attems 1928: 235 (descrip¬

tion)

Sphaerotherium giganteum.— Attems 1934: 465 (descrip¬

tion)

Sphaerotherium giganteum :— Manton 1954: 299 (list)

Sphaerotherium giganteum :— Lawrence 1953: 44 (list)

Sphaerotherium giganteum — Lawrence 1954: 170 (list)

Sphaerotherium (Sphaerotherium) giganteum :— Schubart

1958: 92 (list)

Sphaerotherium giganteum — Lawrence 1987: 109 (des¬

cription)

Sphaerotherium giganteum .— Haacker & Fuchs 1972: 192

(behaviour)

Sphaerotherium (Sphaerotherium) giganteum :— Hamer

1998: 17 (list)

Sphaerotherium giganteum :— Herbert 2000: 2 (prey of

slugs)

Sphaerotherium giganteum — VandenSpiegel et al. 2002:

158 (redescription)

Sphaerotherium giganteum :— Redi et al. 2005: 287 (eco¬

logy)

Sphaerotherium giganteum - Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaerotherium giganteum :—Armstrong & Hamer 2015:

153 (list).

Types: F lectotype, 1 F paralectotype (designated by Van¬

denSpiegel et al. 2002: 158).

Habitat: South Africa: ‘Cafraria’.

Deposited: SMNH.

Sphaerotherium granulatum Pocock, 1895

Sphaerotherium granulatum Pocock, 1895: 410 (first de¬

scription)

Sphaerotherium granulatum :— Silvestri 1910: 196 (de¬

scription)

Sphaerotherium granulatum :—Attems 1928: 217 (descrip¬

tion)

Sphaerotherium (Tetraconosoma) granulatum :— Schubart

1958: 50 (list)

Sphaerotherium (Tetraconosoma) granulatum — Hamer

1998: 17 (list)

Sphaerotherium granulatum :— VandenSpiegel et al. 2002:

176 (list).

Types: Syntypes of unknown number and sex.

Habitat: South Africa: E. Cape: Port Elizabeth.

Deposited: NHML (Online Catalogue).

Sphaerotherium hanstroemi Schubart, 1958

Sphaerotherium (Tetraconosoma) hanstromi [sic!]

Schubart, 1958: 62 (first description)

Sphaerotherium {Tetraconosoma) hanstromi [sic!]:—

Hamer 1998: 17 (list)

Sphaerotherium hanstroemi '.— VandenSpiegel et al.

2002: 176 (list)

Sphaerotherium hanstroemi :— Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium hanstromi [sic!]:— Armstrong & Hamer

2015: 153 (list).

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Species Catalogue Sphaerotheriida

Types: M holotype, 4 F paratypes.

Habitat: South Africa: KZN: Pietermaritzburg, Town

Bush.

Deposited: MZLU.

Sphaerotherium intermedium Porat, 1872

Sphaerotherium intermedium Porat, 1872: 8 (first descrip¬

tion)

Sphaerotherium intermedium.— Preudhomme de Borre

1884: 26 (list)

Sphaerotherium intermedium :— Silvestri 1910: 202 (des¬

cription)

Sphaerotherium intermedium : — Attems 1928: 222 (list)

Sphaerotherium (Tetraconosoma) intermedium :— Schub-

art 1958: 50 (list)

Sphaerotherium (Tetraconosoma) intermedium :— Hamer

1998: 17 (list)

Sphaerotherium intermedium :— VandenSpiegel et al.

2002: 160 (redescription).

Types: F lectotype (designated by VandenSpiegel et al.

2002: 160).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: SMNH.

Sphaerotherium kitharistes Attems, 1928

Sphaerotherium kitharistes Attems, 1928: 216 (first de¬

scription)

Sphaerotherium (Tetraconosoma) kitharistes :— Schubart

1958: 78 (list)

Sphaerotherium (Tetraconosoma) kitharistes :— Hamer

1998: 17 (list)

Sphaerotherium kitharistes :— VandenSpiegel et al. 2002:

176 (list).

Types: 1 M, several F syntypes.

Habitat: Mozambique: Macequece.

Deposited: SAMC, NHMW 3954 (Stagl & Schiller, in

prep.).

Sphaerotherium krugeri Lawrence, 1966

Sphaerotherium krugeri Lawrence, 1966: 242 (first de¬

scription)

Sphaerotherium krugeri :— Hamer 1998: 17 (list)

Sphaerotherium krugeri— VandenSpiegel et al. 2002: 176

(list).

Types: Holotypes [sic!] 2 M, 3 F.

Habitat: South Africa: KNP: Gomondwane.

Deposited: NMSA 9315.

Sphaerotherium lichtensteinii Brandt, 1833

Sphaerotherium Lichtensteinii [sic!] Brandt, 1833: 199

(first description)

ZephroniaLichtenteinii [sic!]:-- Gervais 1837: 9 (list)

Zephronia Lichtenteinii— Gervais 1847: 78 (description)

Zephronia Lichtenteinii— Deshayes & Milne Edwards

1838: 47 (list)

Sphaerotherium Lichtensteinii [sic!]:— Brandt 1841a: 361

(description)

Sphaerotherium Lichtensteinii [sic!]:— Brandt 1841b: 177

(description)

Zephronia Lichtenteinii [sic!]:— Lucas 1842: 521 (descrip¬

tion)

Sphaerotherium Lichtensteinii [sic!]:— Koch 1847: 99

(list)

Sphaerotherium Lichtensteinii [sic!]:— Koch 1863: 33

(description)

Sphaerotherium lichtensteinii :— Butler 1873: 174 (list)

Sphaerotherium Lichtensteinii [sic!]:— Preudhomme de

Borre 1884: 27 (list)

Sphaerotherium lichtensteinii :— Moritz & Fischer 1973:

369 (list)

Sphaerotherium lichtensteinii :— Jeekel 2001: 32 (list)

Sphaerotherium lichtensteinii :— Golovatch & Hoffman

2001: 239 (list)

Sphaerotherium lichtensteinii :— VandenSpiegel et al.

2002: 163 (redescription)

NOT Sphaerotherium Lichtensteinii [sic!]:— Tomosvary,

1885: 68 (list, mis identification) and

Sphaerotherium Lichtensteinii [sic!]:— Daday 1889: 141

(list, misidentification).

Types: 1 F lectotype, 1 F paralectotype (designated by

VandenSpiegel et al. 2002: 163).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: lectotype ZISP, paralectotype ZMB 63.

Sphaerotherium mahaium Schubart, 1958

Sphaerotherium (Tetraconosoma) mahaium Schubart,

1958: 54-56 (first description)

Sphaerotherium (Tetraconosoma) mahaium :— Hamer

1998: 17 (list)

Sphaerotherium mahaium — VandenSpiegel et al. 2002:

177 (list)

Sphaerotherium mahaium — Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium mahaium — Armstrong & Hamer 2015:

153 (list).

Types: M holotype, 1 F paratype.

Habitat: South Africa: KZN: Drakensberg, RNNP, Mahai

River, the Cascades.

Deposited: MZLU.

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Thomas Wesener

Sphaerotherium millepunctatum Attems, 1928

Sphaerotherium millepunctatum Attems, 1928: 219 (first

description)

Sphaerotherium millepunctatum — Attems 1934: 265 (de¬

scription)

Sphaerotherium (Tetraconosoma) millepunctatum —

Schubart 1958: 46 (list)

Sphaerotherium (Tetraconosoma) millepunctatum —

Hamer 1998: 17 (list)

Sphaerotherium millepunctatum :-- VandenSpiegel et al.

2002: 177 (list)

Sphaerotherium millepunctatum :-- Armstrong & Hamer

2015: 153 (list).

Types: 1 M, 1 F syntypes.

Habitat: Mozambique: Masiene near Chai Chai.

Deposited: SAMC, NHMW 2201 (Stagl & Schiller, in

prep.).

Sphaerotherium mo destum Attems, 1928

Sphaerotherium modestum Attems, 1928: 230 (first de¬

scription)

Sphaerotherium (Sphaerotherium) modestum :— Schubart

1958: 93 (list)

Sphaerotherium modestum :— Lawrence 1966: 240 (list)

Sphaerotherium (Sphaerotherium) modestum :— Hamer

1998: 17 (list)

Sphaerotherium modestum :— VandenSpiegel et al. 2002:

177 (list)

Sphaerotherium modestum :— Druce et al. 2007: 211 (ecol¬

ogy)

Sphaerotherium modestum — Druce et al. 2014a: 318 (list)

Sphaerotherium modestum :— Druce et al. 2014b: 297

(list).

Types: F holotype.

Habitat: South Africa: LP: Soutpansberg Distr., 20 miles

E Pietersburg.

Deposited: NHMW 72/230 (Stagl & Schiller, in prep.).

Sphaerotherium narcissei Alderweireldt, 1997

Sphaerotherium narcissei Alderweireldt, 1997: 53 (first

description)

Sphaerotherium narcissei :— Hamer 1998: 18 (list)

Sphaerotherium narcissei :— VandenSpiegel et al. 2002:

177 (list).

Types: M holotype, 4 paratypes of unknown sex.

Habitat: Zimbabwe: Melsetter, 64 km N Chipinga.

Deposited: MRAC.

Sphaerotherium nigritarse Schubart, 1958

Sphaerotherium (Tetraconosoma) nigritarse Schubart,

1958: 81 (first description)

Sphaerotherium (Tetraconosoma) nigritarse :— Hamer

1998: 18 (list)

Sphaerotherium nigritarse :— VandenSpiegel et al. 2002:

177 (list).

Types: F holotype.

Habitat: South Africa: W. Cape: Hermanus.

Deposited: MZLU.

Sphaerotherium obtusum Koch, 1847

Sphaerotherium obtusum Koch, 1847: 98 (first descrip¬

tion)

Sphaerotherium obtusum :— Koch 1863: 5 (description)

Sphaerotherium obtusum :— Preudhomme de Borre 1884:

27 (list)

Sphaerotherium obtusum — Bourne 1885: 162 (descrip¬

tion)

Bournellum obtusum :— de Saussure & Zehntner 1902: 21

(description)

Sphaerotherium obtusum :— Moritz & Fischer 1973: 369

(list)

Sphaerotherium obtusum — VandenSpiegel et al. 2002:

164 (redescription)

Sphaerotherium obtusum :— Armstrong & Hamer 2015:

153 (list).

— Sphaerotherium grossum Koch, 1847: 98 (first des¬

cription), synonymized by VandenSpiegel et al. 2002:

164

Sphaerotherium grossum — Koch 1863: 5 (description)

Sphaerotherium grossum :— Butler 1873: 176 (list)

Sphaerotherium grossum :— Preudhomme de Borre 1884:

26 (list)

Sphaerotherium grossum :— Moritz & Fischer 1973: 369

(list).

Types: F holotype.

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: ZMB 58.

Sphaerotherium perbrincki Schubart, 1958

Sphaerotherium (Tetraconosoma) perbrincki Schubart,

1958: 51 (first description)

Sphaerotherium (Tetraconosoma) perbrincki :— Hamer

1998: 18 (list)

Sphaerotherium perbrincki :— VandenSpiegel et al. 2002:

177 (list)

Sphaerotherium perbrincki :— Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium perbrincki :— Armstrong & Hamer 2015:

153 (list).

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Species Catalogue Sphaerotheriida

Types: M holotype, 1 F paratype.

Habitat: South Africa: KZN: Drakensberg, RNNP, Gudu

Falls area.

Deposited: MZLU.

Sphaerotherium permodestum Silvestri, 1910

Sphaerotherium permodestum Silvestri, 1910: 214 (first

description)

Sphaerotherium permodestum:— Attems 1928: 23 1 (list)

Sphaerotherium (Sphaerotherium)permodestum.— Schub-

art 1958: 94 (list)

Sphaerotherium (Sphaerotherium) permodestum :-- Hamer

1998: 18 (list)

Sphaerotherium permodestum :-- VandenSpiegel et al.

2002: 177 (list).

Types: F holotype.

Habitat: South Africa: Transvaal, no details.

Deposited: IEPN or ZMA.

Sphaerotherium pinnatum Schubart, 1958

Sphaerotherium (Tetraconosoma) pinnatum Schubart,

1958: 76 (first description)

Sphaerotherium (Tetraconosoma) pinnatum :-- Hamer

1998: 18 (list)

Sphaerotherium pinnatum :-- VandenSpiegel et al. 2002:

177 (list).

Types: F holotype.

Habitat: Not specified.

Deposited: MZLU.

Sphaerotherium plagiarium Silvestri, 1910

Sphaerotherium plagiarium Silvestri, 1910: 193 (first de¬

scription)

Sphaerotherium plagiarium.- Attems 1928: 224 (descrip¬

tion)

Sphaerotherium (Tetraconosoma) plagiarium :— Schubart

1958: 86 (list)

Sphaerotherium (Tetraconosoma) plagiarium :— Hamer

1998: 18 (list)

Sphaerotherium plagiarium :— VandenSpiegel et al. 2002:

178 (list).

Types: F holotype.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium punctatum Brandt, 1833

Sphaerotherium punctatum Brandt, 1833: 199 (first de¬

scription)

Zephronia punctata :— Gervais 1837: 9 (list)

Zephroniapunctata :— Deshayes & Milne Edwards 1838:

47 (list)

Sphaerotherium punctatum - Brandt 1841a: 362 (descrip¬

tion)

Sphaerotherium punctatum :— Brandt 1841b: 179 (descrip¬

tion)

Zephronia punctata — Lucas 1842: 522 (description)

Sphaerotherium punctatum :— Newport 1844: 10 (list)

Zephronia punctata :— Gervais 1847: 81 (description)

Sphaerotherium punctatum :— Koch 1847: 98 (list)

Sphaerotherium punctatum :— Koch 1863: 43 (descrip.)

Sphaerotherium punctatum :— Butler 1873: 174 (descrip.)

Sphaerotherium punctatum :— Preudhomme de Borre

1884: 27 (list)

Sphaerotherium punctatum — Moritz & Fischer 1973: 369

(list)

Sphaerotherium punctatum — Lawrence 1987: 11 (list)

Sphaerotherium punctatum :— Golovatch & Hoffman

2001: 239 (list)

Sphaerotherium punctatum :— VandenSpiegel et al. 2002:

166 (redescription).

— Sphaerotherium kochii Butler, 1873: 177, syn-

onymized with S. punctatum by Pocock 1894: 330.

Sphaerotherium kochii :— Preudhomme de Borre 1884: 27

(list).

Types: 1 F lectotype, 1 F paralectotype (selected by Van¬

denSpiegel et al. 2002: 166).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: ZMB 56.

Sphaerotherium punctulatum Brandt, 1841

Sphaerotherium punctulatum Brandt, 1841a: 362 (first de¬

scription)

Sphaerotherium punctulatum :— Brandt 1841b: 179 (de¬

scription)

Zephronia punctulata — Gervais 1847: 81 (description)

Sphaerotherium punctulatum — Porat 1872: 8 (descrip.)

Sphaerotherium punctulatum — Butler 1873: 173 (descr.)

Sphaerotherium punctulatum :— Preudhomme de Borre

1884: 27 (list)

Sphaerotherium punctulatum :— Silvestri 1910: 216 (de¬

scription)

Natalobelum punctulatum :— Verhoeff 1924: 59 (type of

new genus)

Sphaerotherium punctulatum :— Attems 1928: 234 (de¬

scription)

Sphaerotherium punctulatum — Attems 1934: 465 (de¬

scription)

Sphaerotherium punctulatum — Warren 1934: 382 (de¬

scription)

Sphaerotherium (Sphaerotherium) punctulatum —

Schubart 1958: 91 (description)

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Thomas Wesener

Sphaerotherium punctulatum — Haacker & Fuchs 1972:

192 (behaviour)

Sphaerotherium (Sphaerotherium)punctulatum.— Hamer

1998: 18 (list)

Sphaerotherium punctulatum :-- Jolles & Jolles 2000: 234

(medicine)

Sphaerotherium punctulatum :-- Golovatch & Hoffman

2001: 239 (list)

Sphaerotherium punctulatum.— VandenSpiegel 2002: 171

(redescription)

Sphaerotherium punctulatum — VandenSpiegel et al.

2002: 166 (list)

Sphaerotherium punctulatum — Redi et al. 2005: 287

(ecology)

Sphaerotherium punctulatum :-- Regier et al. 2005: 149

(molecular phylogenetic analysis)

Sphaerotherium punctulatum — Wesener & Vanden¬

Spiegel 2009: 548 (morphological phylogenetic analy¬

sis)

Sphaerotherium punctulatum :— Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium punctulatum :— Armstrong & Hamer

2015: 153 (list).

Types: 1 M lectotype (selected by VandenSpiegel 2002:

171).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: ZISR

Sphaerotherium rotundatum Brandt, 1833

Sphaerotherium rotundatum Brandt, 1833: 198 (first de¬

scription)

Zephronia rotundata :— Gervais 1837: 9 (list)

Zephronia rotundata :— Deshayes & Milne Edwards

1838: 47 (list)

Sphaerotherium rotundatum :— Brandt 1841a: 360 (de¬

scription)

Sphaerotherium rotundatum — Brandt 1841b: 175 (de¬

scription

Zephronia rotundata :— Lucas 1842: 521 (description)

Zephronia rotundata :— Gervais 1847: 77 (description)

Sphaerotherium rotundatum :— Koch 1847: 99 (list)

Sphaerotherium rotundatum — Koch 1863: 44(descripti-

on)

Sphaerotherium rotundatum :— Preudhomme de Borre

1884: 28 (list)

Sphaerotherium rotundatum :— Silvestri 1910: 184 (des¬

cription)

Sphaerotherium rotundatum.— Attems 1928: 214 (descrip¬

tion)

Sphaerotherium rotundatum - Attems 1934: 464 (descrip¬

tion)

Sphaerotherium (Tetraconosoma) rotundatum :— Schubart

1958: 75 (list);

Sphaerotherium rotundatum — Moritz & Fischer 1973:

370 (list)

Sphaerotherium (Tetraconosoma) rotundatum :— Hamer

1998: 18 (list)

Sphaerotherium rotundatum :— Golovatch & Hoffman

2001: 239 (list)

Sphaerotherium rotundatum :— VandenSpiegel et al. 2002:

168 (redescription)

Sphaerotherium rotundatum — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaerotherium rotundatum :— Wesener et al. 2011: 967

(stridulation behaviour)

Sphaerotherium rotundatum — Armstrong & Hamer 2015:

153 (list).

— Oligaspis puncticeps Wood 1865: 73 (first descrip¬

tion), synonymized by VandenSpiegel et al. 2002: 168.

Oligaspis puncticeps :— Preudhomme de Borre 1884: 26

(list)

— Sphaerotherium pubescens Porat, 1872: 7 (first de¬

scription), synonymized by Silvestri 1910: 184.

Sphaerotherium pubescens :— Preudhomme de Borre

1884: 27 (list)

— Sphaerotherium viride Porat, 1872: 6 (first descrip¬

tion), synonymized by Silvestri 1910: 184.

Sphaerotherium viride — Preudhomme de Borre 1884: 28

(list).

Types: F holotype.

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: ZMB 61.

Sphaerotherium rudebecki Schubart, 1958

Sphaerotherium (Sphaerotherium) rudebecki Schubart,

1958: 88 (first description)

Sphaerotherium (Sphaerotherium) rudebecki :— Hamer

1998: 18 (list)

Sphaerotherium rudebecki :— VandenSpiegel et al.

2002: 178 (list)

Sphaerotherium rudebecki — Armstrong & Hamer

2015: 153 (list).

Types: M holotype, 1 F paratype.

Habitat: Lesotho: Quthing.

Deposited: MZLU.

Sphaerotherium rugulosum Brandt, 1841

Sphaerotherium rugulosum Brandt, 1841a: 363 (first de¬

scription)

Sphaerotherium rugulosum :— Brandt 1841b: 179 (descrip¬

tion)

Zephronia rugulosa :— Gervais 1847: 82 (description)

Sphaerotherium rugulosum :— Butler 1873: 175 (descrip¬

tion)

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Species Catalogue Sphaerotheriida

Sphaerotherium rugulosum :-- Preudhomme de Borre

1884: 28 (list)

Sphaerotherium rugulosum.— Golovatch & Hoffman

2001:239 (list)

Sphaerotherium rugulosum :-- VandenSpiegel et al. 2002:

171 (redescription).

Remarks: Butler (1873) and VandenSpiegel et al. (2002)

note a potential synonymy of S. rugulosum and S. punc-

tatum.

Types: F lectotype (fragment) (designated by Vanden¬

Spiegel et al. 2002: 171).

Habitat: South Africa: ‘Caput Bonae Spei’ (Cape).

Deposited: ZISP.

Sphaerotherium selindum Alderweireldt, 1997

Sphaerotherium (Tetraconosoma) selindum Alderweireldt,

1997: 53 (first description)

Sphaerotherium selindum :-- Hunt 2002: 803 (parasitic Ne-

matoda)

Sphaerotherium selindum - VandenSpiegel et al. 2002:

178 (list).

Types: M holotype, 1 M, 22 F paratypes.

Habitat: Zimbabwe: Mount Selinda, Chirinda Forest.

Deposited: MRAC.

Sphaerotherium similare Schubart, 1958

Sphaerotherium (Tetraconosoma) similare Schubart,

1958: 82 (first description)

Sphaerotherium (Tetraconosoma) similare — Hamer 1998:

18 (list)

Sphaerotherium similare - VandenSpiegel et al. 2002: 178

(list)

Sphaerotherium similare — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaerotherium similare :-- Wesener et al. 2011: 967

(stridulation behaviour).

Types: M holotype, 1 M, 2 F paratypes.

Habitat: South Africa: E. Cape: Tsitsikama Forest.

Deposited: MZLU.

Sphaerotherium solitarium Attems, 1928

Sphaerotherium solitarium Attems, 1928: 233 (first de¬

scription)

Sphaerotherium (Sphaerotherium) solitarium — Schubart

1958: 91 (list)

Sphaerotherium solitarium :— Lawrence 1966: 241 (list)

Sphaerotherium (Sphaerotherium) solitarium :— Hamer

1998: 19 (list)

Sphaerotherium solitarium — VandenSpiegel et al. 2002:

178 (list).

Types: F holotype.

Habitat: South Africa: LP: Shilowane near Leydsdorp.

Deposited: SAMC; NHMW 72/233 (Stagl & Schiller, in

prep.).

Sphaerotherium spinatum Silvestri, 1910

Sphaerotherium spinatum Silvestri, 1910: 187 (first de¬

scription)

Sphaerotherium spinatum — Attems 1928: 213 (descrip¬

tion)

Sphaerotherium (Tetraconosoma) spinatum :— Schubart

1958: 60 (list)

Sphaerotherium (Tetraconosoma) spinatum :— Hamer

1998: 19 (list)

Sphaerotherium spinatum — VandenSpiegel et al. 2002:

179 (list).

Types: M and F syntypes.

Habitat: South Africa: E. Cape: Port Elizabeth.

Deposited: IEPN or ZMUH.

Sphaerotherium steppense Schubart, 1958

Sphaerotherium (Tetraconosoma) steppense Schubart,

1958: 65 (first description)

Sphaerotherium (Tetraconosoma) steppense :— Hamer

1998: 19 (list)

Sphaerotherium steppense — VandenSpiegel et al. 2002:

179 (list).

Types: M holotype.

Habitat: South Africa: E. Cape: 15 miles S Middleton.

Deposited: MZLU.

Sphaerotherium subdorsale Silvestri, 1910

Sphaerotherium subdorsale Silvestri, 1910: 208 (first de¬

scription)

Sphaerotherium subdorsale - Attems 1928: 228 (descrip.)

Sphaerotherium (Sphaerotherium) subdorsale :— Schubart

1958: 95 (description)

Sphaerotherium (Sphaerotherium) subdorsale :— Hamer

1998: 19 (list)

Sphaerotherium subdorsale :— VandenSpiegel et al. 2002:

179 (list)

Sphaerotherium subdorsale — Armstrong & Hamer 2015:

153 (list).

Types: M and F syntypes.

Habitat: South Africa: Transvaal, no details.

Deposited: IEPN or ZMA.

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Thomas Wesener

Sphaerotherium submite Silvestri, 1910

Sphaerotherium submite Silvestri, 1910: 204 (first descrip¬

tion)

Sphaerotherium submite.— Attems 1928: 222 (list)

Sphaerotherium (Tetraconosoma) submite.— Schubart

1958: 58 (list)

Sphaerotherium (Tetraconosoma) submite:— Hamer 1998:

19 (list)

Sphaerotherium submite :— VandenSpiegel et al. 2002: 179

(list).

Types: M and F syntypes.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium tenuitarse Silvestri, 1910

Sphaerotherium tenuitarse Silvestri, 19 10: 186 (first de¬

scription)

Sphaerotherium tenuitarse:— Attems 1928: 215 (descrip¬

tion)

Sphaerotherium tenuitarse:— Attems 1934: 464 (descrip.)

Sphaerotherium (Tetraconosoma) tenuitarse :— Schubart

1958: 76 (list)

Sphaerotherium (Tetraconosoma) tenuitarse:— Hamer

1998: 19 (list)

Sphaerotherium tenuitarse :— VandenSpiegel et al. 2002:

179 (list)

Sphaerotherium tenuitarse:— Armstrong & Hamer 2015:

153 (list).

Types: M and F syntypes.

Habitat: South Africa: W. Cape: Knysna.

Deposited: IEPN.

Sphaerotherium tomentosum Schubart, 1958

Sphaerotherium (Sphaerotherium) tomentosum Schubart,

1958: 96 (first description)

Sphaerotherium (Sphaerotherium) tomentosum :— Hamer

1998: 19 (list)

Sphaerotherium tomentosum:— VandenSpiegel et al.

2002: 179 (list)

Sphaerotherium tomentosum :— Armstrong & Hamer

2015: 153 (list).

Types: M holotype, 1 F paratype.

Habitat: South Africa: KZN: Drakensberg, RNNP, Gudu

Falls.

Deposited: MZLU.

Sphaerotherium transzambeziacum Jocque, 1984

Sphaerotherium (Tetraconosoma) transzambeziacum

Jocque, 1984: 67 (first description)

Sphaerotherium transzambeziacum :— VandenSpiegel et al.

2002: 179 (list).

Types: M holotype, 12 F paratypes.

Habitat: Malawi, Mt. Mulanje, Tchuchila, Nambiti.

Deposited: Holotype MRAC 15420, paratypes MRAC

14922 & 14938.

Sphaerotherium tuberosum Attems, 1928

Sphaerotherium tuberosum Attems, 1928: 220 (first de¬

scription)

Sphaerotherium (Tetraconosoma) tuberosum :— Schubart

1958: 56 (description)

Sphaerotherium (Tetraconosoma) tuberosum:— Hamer

1998: 19 (list)

Sphaerotherium tuberosum :— VandenSpiegel et al. 2002:

179 (list).

Types: M and F syntypes.

Habitat: South Africa: Cape Province; W. Cape, Table

Mtn; Zoekmakaar, N. Transvaal. No type locality has been

selected.

Deposited: SAMC; NHMW2203.

Sphaerotherium tzitzikamum Schubart, 1958

Sphaerotherium (Tetraconosoma) tzitzikamum Schubart,

1958: 70 (first description)

Sphaerotherium (Tetraconosoma) tzitzikamum :— Hamer

1998: 19 (list)

Sphaerotherium (Tetraconosoma) tzitzikamum:— Van¬

denSpiegel et al. 2002: 179 (list).

Types: M holotype, 1 F paratype.

Habitat: South Africa: E. Cape: Tsitsikama Forest,

Stormsriver.

Deposited: MZLU.

Sphaerotherium weberii Silvestri, 1910

Sphaerotherium Weberii [sic!] Silvestri, 1910: 194 (first

description)

Sphaerotherium weberi [sic!]:— Attems 1928: 219 (de¬

scription)

Sphaerotherium (Tetraconosoma) weberii:— Schubart,

1958: 43 (description)

Sphaerotherium (Tetraconosoma) weberii afoveatum :—

Schubart, 1958: 46 (first description)

Sphaerotherium (Tetraconosoma) weberii :— Hamer 1998:

19 (list)

Sphaerotherium (Tetraconosoma) weberii afoveatum:—

Hamer 1998: 20 (list)

Sphaerotherium (Tetraconosoma) weberii :— VandenSpie¬

gel et al. 2002: 179 (list)

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Species Catalogue Sphaerotheriida

Sphaerotherium (Tetraconosoma) weberii afoveatum —

VandenSpiegel et al. 2002: 179 (list).

Types: M and F syntypes.

Habitat: South Africa: W. Cape: Table Mtn.

Deposited: IEPN.

Sphaerotherium zuluense Schubart, 1958

Sphaerotherium (Tetraconosoma) zuluense Schubart,

1958: 74 (first description)

Sphaerotherium (Tetraconosoma) zuluense.— Hamer

1998: 20 (list)

Sphaerotherium zuluense — VandenSpiegel et al. 2002:

180 (list)

Sphaerotherium zuluense — Armstrong & Hamer 2015:

153 (list).

Types: F holotype.

Habitat: South Africa: KZN: Hluhluwe Game Reserve.

Deposited: MZLU.

Genus Kvlindotherium Attems. 1926

Kylindotherium Attems, 1926: 120 (first description); At¬

tems 1928: 257 (description); Jeekel 1971: 26 (list); Jeekel

1974: 45 (classification); Wesener & VandenSpiegel 2009:

551 (classification); Wesener et al. 2010b: 1188 (classifi¬

cation).

Description: Most likely a synonym of Sphaerotherium.

Type species: Kylindotherium leve Attems, 1926: 118 by

monotypy.

Kylindotherium leve Attems, 1926

Kylindotherium leve Attems, 1926: 118 (first description)

Kylindotherium leve — Attems 1928: 237 (description).

Types: M and F syntypes.

Habitat: South Africa, Cape, Wellington, Witte River.

Deposited: NHMW 2193 (Stagl & Schiller, in prep.).

Nomina dubia of the Sphaerotheriidae

Sphaerotherium nigrum Butler, 1872

Sphaerotherium nigrum Butler 1872: 359 (first descrip¬

tion)

Sphaerotherium nigrum :— Butler 1873: 173 (list)

Sphaerotherium nigrum — Preudhomme de Borre 1884:

27 (list)

Sphaerotherium nigrum — Attems 1928: 236 (list)

Sphaerotherium nigrum - Schubart 1958: 101 (list)

Sphaerotherium nigrum — VandenSpiegel et al. 2002: 180

(list).

Types: 1 holotype of unknown sex.

Habitat: South Africa.

Deposited: NHML (presumably lost).

Family Arthrosphaeridae Jeekel. 1974

Arthrosphaeridae Wesener & VandenSpiegel, 2009: 561.

Arthrosphaeridae Wesener et al. 2010b: 1186 (mentioned

in Sphaerotheriida classification).

Sub Subfamily Arthrosphaerinae Jeekel, 1974: 45 (first de¬

scription); Hoffman 1980: 63 (list); Shelley et al. 2000:

12 (list); Jeekel 2001: 6 (list); Arthrosphaerinae Wesener

& Sierwald 2005b: 598 (cladogram).

Genera included (4)

Arthrosphaera Pocock, 1895 (Southern India and Sri Lan¬

ka)

Sphaeromimus de Saussure & Zehntner, 1902 (SE Mada¬

gascar)

Microsphaerotherium Wesener & VandenSpiegel, 2007

(central Madagascar)

Zoosphaerium Pocock, 1895 (Madagascar)

Type genus: Arthrosphaera Pocock, 1895

117 species (including 11 nomina dubia).

Genus Arthrosphaera Pocock. 1895

(Figs 12 D-E)

Arthrosphaera Pocock, 1895: 410 (first description);

Pocock 1899: 270 (key); Attems 1914: 142 (list); Verho-

eff 1924: 60; Attems 1926: 120 (list); Attems 1928: 203

(new species); Attems 1935: 127 (new species); Attems

1936: 152 (key); Attems 1943: 62 (new species); Attems

1953: 155 (list); Jeekel 1971: 23 (list); Jeekel 1974: 45

(classification); Hoffman 1980: 63 (list); Jeekel 2001: 7

(list); Ashwini & Sridhar 2008: 749 (distribution); Wesen¬

er & VandenSpiegel 2009: 551 (morphological phyloge¬

netic analysis); Wesener et al. 2010b: 1188 (molecular

phylogenetic analysis); Wesener 2014a: 199 (morpholog¬

ical phylogenetic analysis).

Synonyms:

— Trochosoma Chamberlin, 1921: 56 (first description),

synonymized under Arthrosphaera by Jeekel 2001: 7.

Jeekel 1971: 29 (list); Jeekel 1974: 45 (classification);

Hoffman 1980: 63 (list).

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

Type species : Trochosoma pelloceps Chamberlin, 1921

— Trivandrobelum Verhoeff, 1937: 82 (first description),

synonymized under Arthrosphaera by Jeekel 2001: 7.

Verhoeff 1937: 82 (list); Jeekel 1971: 29 (list); Jeekel

1974: 45 (classification); Hoffman 1980: 63 (list).

Type species : Trivandrobelum stridulum Verhoeff 1937

— Odontosternum Attems, 1943: 62 (first description),

synonymized under Arthrosphaera by Jeekel 2001: 7.

Attems 1953: 155 (list); Jeekel 1971: 27 (list); Jeekel

1974: 45 (classification).

Type species : Odontosternum fuscum Attems, 1943

Description: Anterior telopod with four podomeres dis¬

tal to syncoxite, telopoditomere 4 with lobe covered with

sclerotized spots. Harp carrying one or two stridulation

ribs. Posterior telopods with three telopoditomeres,

telopoditomere 3 carrying two membranous lobes. Oper¬

culum of female vulva well-rounded, shorter than bursa.

Inner plate of vulva heavily elongated, protruding as far

as, or even above, operculum. Female washboard with

few, usually 4-6 stridulation ribs. Legs with a coxal lobe,

tarsi usually wide, paddle-like, rarely slender. Underside

of anal shield usually with two locking carinae, which are

completely reduced in some species. First antennomere

without groove, only isolated sclerotized scales, lacking

sensilla basiconica like all other antennomeres. Anten¬

nomere 6 large, flattened, axe-shaped, carrying numerous

apical cones. 25-45 mm long.

Type species: Zephronia heterostictica Newport, 1844.

Included species (40)

Arthrosphaera atrisparsa (Butler, 1878) (India)

Arthrosphaera attemsi Jeekel, 2001 (Sri Lanka)

Arthrosphaera aurocincta Pococlc, 1899 (India)

Arthrosphaera bicolor Pocock, 1895 (India)

Arthrosphaera brandti (Humbert, 1865) (Sri Lanka, intro¬

duced to Tanzania)

Arthrosphaera carinata Attems, 1936 (India)

Arthrosphaera corrugata (Butler, 1872) (Sri Lanka)

Arthrosphaera craspedota Attems, 1936 (India)

Arthrosphaera dalyi Pocock, 1895 (India)

Arthrosphaera davisoni Pocock, 1895 (India)

Arthrosphaera dentigera Verhoeff, 1930 (Sri Lanka)

Arthrosphaera disticta Pocock, 1895 (India)

Arthrosphaera fumosa Pocock, 1895 (India)

Arthrosphaera fusca (Attems, 1943) (India)

Arthrosphaera gracilis Attems, 1936 (India)

Arthrosphaera hendersoni Pocock, 1895 (India)

Arthrosphaera heterostictica (Newport, 1844) (India)

Arthrosphaera inermis (Humbert, 1865) (India, Sri Lan¬

ka)

Arthrosphaera leopardina (Butler, 1872) (Sri Lanka)

Arthrosphaera lutescens (Butler, 1872) (India)

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera madura Pandian, 1966 (India)

Arthrosphaera magna Attems, 1936 (India)

Arthrosphaera marginella Silvestri, 1897 (India)

Arthrosphaera marmorata (Butler, 1882) (India)

Arthrosphaera nitida Pocock, 1895 (India)

Arthrosphaera noticeps (Butler, 1872) (Sri Lanka)

Arthrosphaera pelloceps (Chamberlin, 1921) (India)

Arthrosphaera pilifera (Butler, 1872) (Sri Lanka)

Arthrosphaera pygostolis Attems, 1935 (India)

Arthrosphaera ruginosa Jeekel, 2001 (Sri Lanka)

Arthrosphaera rugosa Verhoeff, 1930 (Sri Lanka)

Arthrosphaera scholastica Attems, 1936 (India)

Arthrosphaera severa Attems, 1935 (India)

Arthrosphaera silvestrii Jeekel, 2001 (India)

Arthrosphaera stridula (Verhoeff, 1937) (India)

Arthrosphaera thurstoni Pocock, 1895 (India)

Arthrosphaera transitiva Attems, 1936 (India)

Arthrosphaera versicolor (White, 1859) (Sri Lanka)

Arthrosphaera wroughtoni Pocock, 1895 (India)

Arthrosphaera zebraica (Butler, 1872) (India)

Arthrosphaera atrisparsa (Butler, 1878)

Zephronia atrisparsa Butler, 1878: 302 (first description)

Zephronia atrisparsa:— Preudhomme de Borre 1884: 23

(list)

Arthrosphaera atrisparsa :— Pocock 1899: 274 (new lo¬

cality)

Arthrosphaera atrisparsa:— Attems 1914: 147 (list)

Arthrosphaera atrisparsa :— Attems 1936: 164 (list)

Arthrosphaera artisparsa [sic!]:— Rajulu 1970: 125 (list)

Arthrosphaera atrisparsa :— Kunte 1974: 673 (list)

Arthrosphaera atrisparsa :— Jeekel 2001: 7 (list)

Arthrosphaera atrisparsa :— Sridhar & Kadamannaya

2009: 68 (list)

Arthrosphaera atrisparsa :— Sridhar 2015: 3 (list)

Arthrosphaera atrisparsa:— Sridhar &Ashwini 2016: 18

(list).

— Zephronia heterosticatica [sic!] e.p. :— Pocock 1892:

145, NOT Zephronia heterostictica Newport, 1844.

Types: M holotype.

Locality: India: Bombay.

Deposited: NHML.

Arthrosphaera attemsi Jeekel, 2001

Arthrosphaera attemsi Jeekel, 2001: 7 (e.p.)

Arthrosphaera attemsi :— De Zoysa et al. 2016: 454 (list).

— Arthrosphaera inermis Attems 1936: 159 (first de¬

scription, e.p.), NOT Arthrosphaera inermis (Humbert,

1865).

Types: M holotype.

Locality: Sri Lanka: Peradeniya.

Deposited: NHMW 100/159 (Stagl & Schiller, in prep.).

©ZFMK

Species Catalogue Sphaerotheriida

Arthrosphaera aurocincta Pocock, 1899

Arthrosphaera aurocincta Pocock, 1899: 276 (first de¬

scription)

Arthrosphaera aurocincta:— Gravely 1915: 518 (new lo¬

cality, sound)

Arthrosphaera aurocincta:— Attems 1914: 147 (list)

Arthrosphaera aurocincta:— Attems 1936: 165 (list)

Arthrosphaera aurocincta :— Rajulu 1970: 125 (list)

Arthrosphaera aurocincta :— Jeekel 2001: 7 (list)

Arthrosphaera aurocincta :— Sridhar & Kadamannaya

2009: 68 (list)

Arthrosphaera aurocincta :— Sridhar 2015: 3 (list)

Arthrosphaera aurocincta :— Sridhar & Ashwini 2016: 18

(list).

Types: M and F syntypes.

Locality: “East Indies” (Pocock 1899), Parambikulam, In¬

dia: Cochin Ghats (Gravely 1915).

Deposited: NHML.

Arthrosphaera bicolor Pocock, 1895

Arthrosphaera bicolor Pocock, 1895: 411 (first descrip.)

Arthrosphaera bicolor — Pocock 1899: 278 (description)

Arthrosphaera bicolor.— Chamberlin & Wang 1953: 2

(new locality)

Arthrosphaera bicolor.— Attems 1914: 147 (list)

Arthrosphaera bicolor — Attems 1936: 165 (list)

Arthrosphaera bicolor.— Rajulu 1970: 125 (list)

Arthrosphaera bicolor.— Chowdaiah & Kanaka 1974: 56

(cytology)

Arthrosphaera bicolor — Rangaswamy et al. 1978: 99 (dis¬

tribution)

Arthrosphaera bicolor — Bano 1983: 168 (list)

Arthrosphaera bicolor.- Rangaswamy et al. 1983: 7 (dis¬

tribution)

Arthrosphaera bicolor — Jeekel 2001: 8 (list)

Arthrosphaera bicolor.— Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera bicolor — Sridhar 2015: 3 (list)

Arthrosphaera bicolor :— Sridhar & Ashwini 2016: 18

(list).

Types: M holotype.

Habitat: India: Salem, Shevaroy Hills.

Deposited: NHML.

Arthrosphaera brandti (Humbert, 1865)

(Fig. 13 A)

Sphaeropoeus Brandtii Humbert, 1865: 38 (first descrip¬

tion)

Zephronia Brandti — Butler 1872: 355 (list)

Zephronia brandtii :— Butler 1873: 178 (list)

Zephronia Brandtii — Karsch 1881: 29 (list)

Zephronia Brandtii — Preudhomme de Borre 1884: 23

(list)

Zephronia brandti [sic!]:— Pocock 1892: 143 (new local¬

ity)

Arthrosphaera Brandti :— Pocock 1899: 272 (new locali-

ty)

Sphaeropoeus Brandti — Saussure & Zehntner 1902: 14

(list)

Arthrosphaera Brandti:— Attems 1914: 147 (list)

Arthrosphaera brandti :— Carl 1922: 565 (list)

Arthrosphaera brandti :— Attems 1936: 164 (description)

Arthrosphaera brandti :— Rajulu 1970: 125 (list)

Arthrosphaera brandti — Enghoff 1977: 997 (new local¬

ity, introduced E. Africa)

Arthrosphaera brandti :— Jeekel 2001: 8 (list)

Arthrosphaera brandti :— Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera brandti :— Wesener & VandenSpiegel

2009: 548 (phylogenetic analysis)

Arthrosphaera brandti :— Wesener et al. 2010b: 1185 (phy¬

logenetic analysis)

Arthrosphaera brandti :— Pick et al. 2014: 3 (hemocyanin)

Arthrosphaera brandti :— Sridhar 2015: 3 (list)

Arthrosphaera brandti :— Sridhar & Ashwini 2016: 18

(list)

Arthrosphaera brandti :— De Zoysa et al. 2016: 454 (list)

— Zephronia chitinoides Butler, 1872: 354 (first descr.),

synonymized under A. brandtii by Pocock 1892: 143

Zephronia chitonoides [sic!]:— Preudhomme de Borre

1884: 24 (list).

Remark: The record from an East African site concerns

an introduced population (Enghoff 1977).

Types: Syntypes.

Habitat: Sri Lanka: Peradeniya (Type); Punduloya; Ta-

lawa, Anuradhapura. India,Madras. Tanzania: Amani, East

Usambara Mts.

Deposited: MHNG?

Arthrosphaera carinata Attems, 1936

Arthrosphaera carinata Attems, 1936: 162 (first descrip.)

Arthrosphaera carinate [sic!]:— Rajulu 1970: 124 (list)

Arthrosphaera carinata :— Jeekel 2001: 8 (list)

Arthrosphaera carinata :— Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera carinata - Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera carinata :— Sridhar 2015: 3 (list)

Arthrosphaera carinata - Ambarish & Sridhar 2015c (dis¬

tribution)

Arthrosphaera carinata :— Sridhar & Ashwini 2016: 18

(list).

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Thomas Wesener

Types: M syntype.

Habitat: India: Kemmangundi -Kalhaattigiri, Babudan

Hills, Kaddur distr., Mysore.

Deposited: NHMW 2206 (Stagl & Schiller, in prep.).

Arthrosphaera corrugata (Butler, 1872)

Zephronia corrugata Butler, 1872: 355 (first description)

Zephronia corrugata.— Butler 1873: 180 (list)

Zephronia corrugata :— Preudhomme de Borre 1884: 24

(list)

Arthrosphaera corrugata :— Pocock 1899: 273 (new

combination, list)

Arthrosphaera corrugata — Attems 1914: 147 (list);

Arthrosphaera corrugata:— Attems 1936: 157 (new local¬

ity)

Arthrosphaera corrugata :— Jeekel 2001: 8 (list)

Arthrosphaera corrugata — Sridhar & Kadamannaya

2009: 68 (list)

Arthrosphaera corrugata — Sridhar 2015: 3 (list)

Arthrosphaera corrugata — Sridhar & Ashwini 2016: 18

(list)

Arthrosphaera corrugata :— De Zoysa et al. 2016: 454

(list).

— Zephronia inermis (e.p.):— Pocock, 1892: 144. NOT

Sphaeropoeus inermis Humbert, 1865, and NOT

Arthrosphaera corrugata Silvestri, 1897 (=

Arthrosphaera silvestrii Jeekel, 2001).

Types: Holotype of unknown sex.

Habitat: Sri Lanka: Peradeniya.

Deposited: NHML.

Arthrosphaera craspedota Attems, 1936

Arthrosphaera craspedota Attems, 1936: 156

Arthrosphaera craspedota — Rajulu 1970: 124 (list)

Arthrosphaera craspedota — Kanaka & Chowdaiah 1974:

269 (anatomy)

Arthrosphaera craspedota :— Chowdaiah & Kanaka 1974:

56 (cytology)

Arthrosphaera craspedota :— Rangaswamy et al. 1978: 99

(distribution)

Arthrosphaera craspedota :— Bano 1983: 168 (list)

Arthrosphaera craspedota — Rangaswamy et al. 1983: 7

(distribution)

Arthrosphaera craspedota :— Jeekel 2001: 8 (list)

Arthrosphaera craspedata [sic!]:- Sridhar & Kadaman¬

naya 2009: 68 (list)

Arthrosphaera craspedota :— Kadamannaya et al. 2012:

183 (morphology)

Arthrosphaera craspedota :— Ambarish et al. 2013: 205

(cytology)

Arthrosphaera craspedota :— Basil-Rose et al. 2014: 332

(physiology)

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera craspedota — Sridhar 2015: 3 (list)

Arthrosphaera craspedota :— Sridhar & Ashwini 2016: 18

(list).

Types: M and F syntypes.

Habitat: India: Cochin State, Parambikulam.

Deposited: NHMW 2207 (Stagl & Schiller, in prep.).

Arthrosphaera dalyi Pocock, 1895

Arthrosphaera dalyi Pocock, 1895: 412 (first description)

Arthrosphaera dalyi — Pocock 1899: 280 (new locality,

redescription)

Arthrosphaera dalyi — Attems 1914: 147 (list)

Arthrosphaera dalyi:- Attems 1936: 165 (list)

Arthrosphaera dalyi - Rajulu 1970: 125 (list)

Arthrosphaera dalyi — Chowdaiah & Kanaka 1974: 54

(cytology)

Arthrosphaera dalyi — Demange 1977: 231 (new locali¬

ty)

Arthrosphaera dalyi — Rangaswamy et al. 1978: 98 (dis¬

tribution)

Arthrosphaera dalyi - Bano 1983: 168 (list)

Arthrosphaera dalyi — Janardanan & Ramachandran

1983: 408 (parasites)

Arthrosphaera dalyi — Rangaswamy et al. 1983: 7 (dis¬

tribution)

Arthrosphaera dalyii [sic!]: - Nattuthurai & Subbiah 1983:

113 (physiology)

Arthrosphaera dalyi — Achar 1986: 90 (cytology)

Arthrosphaera dalyi - Nandre & Joshi 1993: 40 (morphol¬

ogy)

Arthrosphaera dalyi — Jeekel 2001: 8 (list)

Arthrosphaera dalyi — Sridhar & Kadamannaya 2009: 68

(list)

Arthrosphaera dalyi — Kadamannaya & Sridhar 2009a:

761 (ecology, new locality)

Arthrosphaera dalyi — Kadamannaya & Sridhar 2009b:

506 (behaviour)

Arthrosphaera dalyi — Kadamannaya et al. 2010a: 467

(cytology)

Arthrosphaera dalyi — Kadamannaya et al. 2010b: 245

(ecology)

Arthrosphaera dalyi — Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera dalyi — Sridhar et al. 2013: 539 (ecology)

Arthrosphaera dalyi - Ambarish & Sridhar 2014c: 15 (be¬

haviour)

Arthrosphaera dalyi — Sridhar 2015: 3 (list)

Arthrosphaera dalyi — Ambarish & Sridhar 2015c: (dis¬

tribution)

Arthrosphaera dalyi - Ambarish & Sridhar 2016: 105 (be¬

havior)

Arthrosphaera dalyi — Sridhar & Ashwini 2016: 18 (list).

©ZFMK

Species Catalogue Sphaerotheriida

Types: Syntypes.

Habitat: India: Lone Cottage, Palni Hills; Kodaikanal, Pal-

ni Hills, Western Ghats.

Deposited: NHML.

Arthrosphaera davisoni Pocock, 1895

Arthrosphaera Davisoni Pocock, 1895: 412 (first descrip.)

Arthrosphaera Davisoni.— Pocock 1899: 279 (redescrip¬

tion)

Arthrosphaera davisoni — Attems 1914: 148 (list)

Arthrosphaera davisoni :-- Attems 1936: 165 (list)

Arthrosphaera davisoni.— Rajulu 1970: 125 (list)

Arthrosphaera davisoni.— Bano 1983: 168 (list)

Arthrosphaera davisoni — Achar 1986: 90 (cytology)

Arthrosphaera davisoni — Jeekel 2001: 8 (list)

Arthrosphaera davisoni :-- Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera davisoni — Kadamannaya & Sridhar

2009a: 761 (ecology, new locality)

Arthrosphaera davisoni — Kadamannaya & Sridhar

2009b: 506 (behaviour)

Arthrosphaera davisoni— Kadamannaya et al. 2010b: 245

(ecology)

Arthrosphaera davisoni — Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera davisoni :-- Sridhar et al. 2013: 539 (ecol¬

ogy)

Arthrosphaera davisoni :— Sridhar 2015: 3 (list)

Arthrosphaera davisoni :— Sridhar & Ashwini 2016: 18

(list).

Remark: The spelling “davidsoni” in the 1895 publication

is an obvious printer’s error (Jeekel 2001).

Types: M and F syntypes.

Habitat: India: Coimbatore, Animallai Hills, Western

Ghats.

Deposited: NHML.

Arthrosphaera dentigera Verhoeff, 1930

Arthrosphaera dentigera Verhoeff, 1930: 208 (first de¬

scription)

Arthrosphaera dentigera:— Attems 1936: 165 (list)

Arthrosphaera dentigera :— Jeekel 2001: 9 (list)

Arthrosphaera dentigera — Sridhar & Kadamannaya

2009: 68 (list)

Arthrosphaera dentigera :— Sridhar 2015: 3 (list)

Arthrosphaera dentigera :— Sridhar & Ashwini 2016: 18

(list)

Arthrosphaera dentigera :— De Zoysa et al. 2016: 455 (list)

Remark: The name of this species was already used by

Verhoeff, 1910: 241, fig. XXIX, as a nomen nudum

(Jeekel 2001).

Bonn zoological Bulletin - Supplementum 63: 1-104

Types: M holotype?

Habitat: Sri Lanka: Kandy.

Deposited: Unknown.

Arthrosphaera disticta Pocock, 1895

Arthrosphaera disticta Pocock, 1895: 411 (first descrip¬

tion)

Arthrosphaera disticta :— Pocock 1899: 278 (redescription)

Arthrosphaera disticta :— Attems 1914: 148 (list)

Arthrosphaera disticta :— Attems 1936: 165 (list)

Arthrosphaera disticta :— Rajulu 1970: 125 (list)

Arthrosphaera disticta :— Chowdaiah & Kanaka 1974: 54

(cytology)

Arthrosphaera disticta: :— Demange 1977: 231 (new local¬

ity)

Arthrosphaera disticta— Rangaswamy et al. 1978: 98 (dis¬

tribution)

Arthrosphaera disticta — Bano 1983: 168 (list)

Arthrosphaera disticta — Janardanan & Ramachandran

1983: 408 (parasites)

Arthrosphaera disticta :— Rangaswamy et al. 1983: 6 (dis¬

tribution)

Arthrosphaera disticta :— Mary & Prema 1984: 29 (phys¬

iology)

Arthrosphaera disticta :— Achar 1986: 90 (cytology)

Arthrosphaera disticta :— Jeekel 2001: 9 (list)

Arthrosphaera disticta :— Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera disticta :— Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera disticta :— Ambarish & Sridhar 2014a: 95

(ecology)

Arthrosphaera disticta :— Ambarish & Sridhar 2014c: 15

(behaviour)

Arthrosphaera disticta :— Basil-Rose et al. 2014: 332

(physiology)

Arthrosphaera disticta :— Sridhar 2015: 3 (list)

Arthrosphaera disticta :—Ambarish & Sridhar 2015c: (dis¬

tribution)

Arthrosphaera disticta :— Ambarish & Sridhar 2016: 105

(behaviour).

Arthrosphaera disticta :— Sridhar & Ashwini 2016: 18

(list).

Types: M holotype?

Habitat: India: Yercand, Sheveroy Hills.

Deposited: NHML.

Arthrosphaera fumosa Pocock, 1895

Arthrosphaera fumosa Pocock, 1895: 412 (first descrip¬

tion)

Arthrosphaera fumosa :— Pocock 1899: 280 (redescription)

Arthrosphaera fumosa :— Attems 1914: 148 (list)

©ZFMK

Thomas Wesener

Arthrosphaera fumosa — Attems 1936: 165 (list)

Arthrosphaera fumosa.— Rajulu 1970: 125 (list)

Arthrosphaera fumosa :— Jeekel 2001: 9 (list)

Arthrosphaera fumosa — Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera fumosa :— Kadamannaya & Sridhar 2009b:

506 (behaviour)

Arthrosphaera fumosa — Kadamannaya et al. 2010b: 245

(ecology)

Arthrosphaera fumosa :— Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera fumosa — Ambarish & Sridhar 2013b: 258

(ecology)

Arthrosphaera fumosa :— Sridhar & Ambarish 2013: 1544

(ecology)

Arthrosphaera fumosa — Ambarish & Sridhar 2014a: 95

(ecology)

Arthrosphaera fumosa :— Ambarish & Sridhar 2014b: 49

(cytology)

Arthrosphaera fumosa — Sridhar et al. 2013: 539 (ecolo¬

gy)

Arthrosphaera fumosa :— Ambarish & Sridhar 2014c: 15

(behaviour)

Arthrosphaera fumosa — Sridhar 2015: 3 (list)

Arthrosphaera fumosa— Ambarish & Sridhar 2015a: 642

(ecology)

Arthrosphaera fumosa- Ambarish & Sridhar 2015b: 118

(physiology)

Arthrosphaera fumosa:— Ambarish & Sridhar 2015c: (dis¬

tribution)

Arthrosphaera fumosa — Ambarish & Sridhar 2016: 105

(behavior)

Arthrosphaera fumosa :— Sridhar & Ashwini 2016: 18

(list).

Types: F holotype.

Habitat: India: Coimbatore.

Deposited: NHML.

Arthrosphaera fusca (Attems, 1943)

Odontosternum fuscum Attems, 1943: 65 (first descrip¬

tion)

Arthrosphaera fusca- Jeekel 2001: 9 (new combination).

Types: M and F syntypes.

Habitat: India: Ahmednagar.

Deposited: NHMW 2205 (Stagl & Schiller, in prep.).

Arthrosphaera gracilis Attems, 1936

Arthrosphaera gracilis Attems, 1936: 158 (first descrip.)

Arthrosphaera gracilis :— Rajulu 1970: 124 (list)

Arthrosphaera gracilis :— Chowdaiah & Kanaka 1974: 54

(cytology)

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera gracilis :— Rangaswamy et al. 1978: 98

(distribution)

Arthrosphaera gracilis — Bano 1983: 168 (list)

Arthrosphaera gracilis :— Rangaswamy et al. 1983: 7 (dis¬

tribution)

Arthrosphaera gracilis :— Jeekel 2001: 9 (list)

Arthrosphaera gracilis :— Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera gracilis :— Sridhar 2015: 3 (list)

Arthrosphaera gracilis — Sridhar & Ashwini 2016: 18

(list).

Types: M syntype.

Habitat: India: Courlallam, South India.

Deposited: NHMW 2208 (Stagl & Schiller, in prep.).

Arthrosphaera hendersoni Pocock, 1895

Arthrosphaera Headersoni Pocock, 1895: 411 (first de¬

scription)

Arthrosphaera Hendersoni :— Pocock 1899: 277(re-

description)

Arthrosphaera hendersoni — Attems 1914: 148 (list)

Arthrosphaera hendersoni :— Attems 1936: 162 (addition¬

al specimens)

Arthrosphaera hendersoni :— Rajulu 1970: 124 (list)

Arthrosphaera hendersoni :— Chowdaiah & Kanaka 1974:

54 (cytology)

Arthrosphaera handersoni [sic!]:— Rangaswamy et al.

1978: 99 (distribution)

Arthrosphaera hendersoni :— Bano 1983: 168 (list)

Arthrosphaera hendersoni :— Rangaswamy et al. 1983: 7

(distribution)

Arthrosphaera hendersoni :— Jeekel 2001: 9 (list)

Arthrosphaera hendersoni — Beron 2008: 175 (list)

Arthrosphaera hendersoni — Sridhar & Kadamannaya

2009: 68 (list)

Arthrosphaera hendersoni :— Sridhar 2015: 3 (list)

Arthrosphaera hendersoni :— Ambarish & Sridhar 2015c:

(distribution)

Arthrosphaera hendersoni : — Sridhar & Ashwini 2016: 18

(list).

Types: M and F syntypes.

Habitat: India: Kodaikanal, Palnai Hills.

Deposited: NHML.

Arthrosphaera heterostictica (Newport, 1844)

Zephronia heterostictica Newport, 1844: 265 (first de¬

scription)

Zephronia heterostictica :— Gervais 1847: 84 (list)

Zephronia heterosticta [sic!]:— Preudhomme de Borre

1884: 24 (list)

Zephronia heterostictica — Pocock 1892: 145 (description)

©ZFMK

Species Catalogue Sphaerotheriida

Arthrosphaeraheterostictica — Pocock 1895: 411 (list)

Arthrosphaera heterosticta [sic!]:— Pocock 1899: 273

(new locality)

Arthrosphaera heterosticta [sic!]:— Attems 1936: 165

(new localities)

Arthrosphaera heterosticta [sic!]:— Rajulu 1970: 126 (list)

Arthrosphaera heterostictica :— Jeekel 2001: 9 (list)

Arthrosphaera heterosticta [sic!]:— Sridhar & Kadaman-

naya 2009: 68 (list)

Arthrosphaera heterosticta [sic!]:— Sridhar 2015: 3 (list)

Arthrosphaera heterosticta [sic!]:— Sridhar & Ashwini

2016: 18 (list).

Types: 1 holotype of unknown sex.

Habitat: India: Bombay; Madras; Nilghiri Hills; Panch-

gani, Western Ghats; Poona, Bombay Presidency.

Deposited: NHML.

Arthrosphaera inermis (Humbert, 1865)

Sphaeropoeus inermis Humbert, 1865: 39 (first descrip.)

Zephronia inermis.— Butler 1873: 179 (new locality)

Zephronia inermis — Karsch 1881: 29 (description)

Zephronia inermis — Preudhomme de Borre 1884: 24 (list)

Zephronia inermis — Pocock 1892: 144 (new locality)

Arthrosphaera inermis — Pocock 1899: 273 (new locali¬

ty)

Sphaeropoeus inermis — Saussure & Zehntner 1902: 14

(list)

Arthrosphaera inermis — Attems 1914: 148 (list)

Arthrosphaera inermis :— Jeekel 2001: 9 (list)

Arthrosphaera inermis — Sridhar & Kadamannaya 2009:

68 (list)

Arthrosphaera inermis — Wesener & VandenSpiegel

2009: 548 (phylogenetic analysis)

Arthrosphaera inermis — Sridhar 2015: 3 (list)

Arthrosphaera inermis — Sridhar & Ashwini 2016: 18

(list)

Arthrosphaera inermis :— De Zoysa et al. 2016: 455 (list).

Remark: Not Arthrosphaera inermis Attems, 1936 =

Arthrosphaera attemsi Jeekel, 2001.

Types: syntypes

Habitat: Sri Lanka: Peradeniya; valley of Pundeloya, near

Rambodde; Pundeloya. India: Madras.

Deposited: MHNG.

Arthrosphaera leopardina (Butler, 1872)

Zephronia leopardina Butler, 1872: 356 (first description)

Zephronia leopardina — Butler 1873: 181 (description)

Zephronia leopardina — Preudhomme de Borre: 24 (list)

Arthrosphaera leopardina :— Pocock 1899: 274 (new com¬

bination)

Arthrosphaera leopardina :— Attems 1914: 148 (list)

Arthrosphaera leopardina :— Attems 1936: 166 (list)

Arthrosphaera leopardina :— Jeekel 2001: 10 (list)

Arthrosphaera leopardina — Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera leopardina :— Sridhar 2015: 3 (list)

Arthrosphaera leopardina — Sridhar & Ashwini 2016: 18

(list)

Arthrosphaera leopardina :— De Zoysa et al. 2016: 455

(list).

— Zephronia inermis Pocock 1892: 144 (misidentifica¬

tion, e. p.). NOT Sphaeropoeus inermis Humbert, 1865.

Types: Immature holotype of unknown sex.

Habitat: Sri Lanka.

Deposited: NHML.

Arthrosphaera lutescens (Butler, 1872)

Zephronia lutescens Butler, 1872: 356 (first description)

Zephronia lutescens — Butler 1873: 179 (description)

Zephronia lutescens — Preudhomme de Borre: 25 (list)

Arthrosphaera lutescens :— Pocock 1899: 274 (new com¬

bination)

Arthrosphaera lutescens :— Attems 1914: 148 (list)

Arthrosphaera lutescens :— Attems 1936: 155 (new local¬

ity)

Arthrosphera [sic!] lutescens :— Gowri & Sethuramakrish-

nan 1980: 113 (hormones)

Arthrosphaera lutescens — Sarasu & Sivagamasundari

1982: 74 (eggs)

Arthrosphaera lutescens .— Bano 1983: 168 (list)

Arthrosphaera lutescens — Rangaswamy et al. 1983: 7

(distribution)

Arthrosphaera lutescens :— Achar 1986: 90 (cytology)

Arthrosphaera lutescens :— Jeekel 2001: 10 (list)

Arthrosphaera lutescens :— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera lutescens — Kadamannaya etal. 2012: 183

(morphology)

Arthrosphaera lutescens — Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera lutescens — Ambarish et al. 2013: 205 (cy¬

tology)

Arthrosphaera lutescens :— Sridhar 2015: 3 (list)

Arthrosphaera lutescens :— Sridhar & Ashwini 2016: 18

(list)

— Zephronia heterosticatica [sic!] (misidentification,

e.p.) Pocock, 1892: 145. NOT Zephronia heterosticti¬

ca Newport, 1844.

Remark: According to Pocock (1892) potentially a syn¬

onym of A. heterosticta.

Types: Holotype of unknown sex.

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

Habitat: India: Maddathoray; Kulattupuzha, Western

Ghats, Travancore.

Details: NHML.

Arthrosphaera madura Pandian, 1966

Arthrosphaera madura Pandian, 1966: 465 (first descrip.)

Arthrosphaera madura.— Rajulu 1970: 126 (list)

Arthrosphaera madura :-- Rangaswamy et al. 1983: 6 (dis¬

tribution).

Types: Unknown number of male and female type spec¬

imens.

Habitat: India, Madurai, Algarkoil hills, Pazhamudir So-

lai.

Deposited: Indian Museum Calcutta (Pandian 1966:

468), now Zoological Survey of India, Kolkata.

Arthrosphaera magna Attems, 1936

Arthrosphaera magna Attems, 1936: 160 (first descrip¬

tion)

Arthrosphaera magna — Rajulu 1970: 124 (list)

Arthrosphaera magna — Bano 1983: 168 (list)

Arthrosphaera magna:— Achar 1986: 90 (cytology)

Arthrosphaera magna:— Mary Bai 1997: 461 (list)

Arthrosphaera magna — Jeelcel 2001: 10 (list)

Arthrosphaera magna - Ashwini & Sridhar 2002: 20

(ecology)

Arthrosphaera magna — Ashwini & Sridhar 2005: 307

(ecology)

Arthrosphaera magna — Ashwini & Sridhar 2006a: 954

(ecology)

Arthrosphaera magna — Ashwini & Sridhar 2006b: 27

(ecology)

Arthrosphaera magna — Ashwini & Sridhar 2006c: 59

(ecology)

Arthrosphaera magna — Ashwini & Sridhar 2008: 749

(distribution)

Arthrosphaera magna — Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera magna — Kadamannaya et al. 2010b: 245

(ecology)

Arthrosphaera magna — Kampfer et al. 2010: 1765 (bac¬

teria)

Arthrosphaera magna:— Kampfer et al. 2011: 1334 (bac¬

teria)

Arthrosphaera magna — Ramanthan & Alagesan 2011:

365 (ecology)

Arthrosphaera magna — Ramanthan & Alagesan 2012: 57

(bacteria)

Arthrosphaera magna — Alagesan & Ramanathan 2013:

2 (distribution)

Arthrosphaera magna:— Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera magna — Ambarish & Sridhar 2013b: 258

(ecology)

Arthrosphaera magna — Sridhar & Ambarish 2013: 1544

(ecology)

Arthrosphaera magna:— Sridhar et al. 2013: 539 (ecolo¬

gy)

Arthrosphaera magna — Ambarish & Sridhar 2014b: 49

(cytology)

Arthrosphaera magna:— Ambarish & Sridhar 2014c: 15

(behaviour)

Arthrosphaera magna:— Sridhar 2015: 3 (list)

Arthrosphaera magna — Ambarish & Sridhar 2015a: 642

(ecology)

Arthrosphaera magna:— Ambarish & Sridhar 2015b: 118

(physiology)

Arthrosphaera magna:— Ambarish & Sridhar 2015c: (dis¬

tribution)

Arthrosphaera magna:— Ambarish & Sridhar 2016: 105

(behavior)

Arthrosphaera magna:— Sridhar & Ashwini 2016: 19

(list).

Types: M and F syntypes.

Habitat: India: Khandala, Lonavla, Bombay Presidency;

Shevaroy Hills, S. India; Rajamandri, Madras Presiden¬

cy; Phenda Ghat, Kolhapur State, Bombay Presidency.

Deposited: NHMW 2210 (Stagl & Schiller, in prep.).

Arthrosphaera marginella Silvestri, 1897

Arthrosphaera marginella Silvestri, 1897b: 360 (first de¬

scription)

Arthrosphaera marginella :— Pocock 1899: 285;

Arthrosphaera marginella:— Attems 1914: 148 (list);

Arthrosphaera marginella — Attems 1936: 166 (list)

Arthrosphaera marginella — Jeekel 2001: 10 (list)

Arthrosphaera marginella :— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera marginella - Sridhar 2015: 3 (list)

Arthrosphaera marginella — Sridhar & Ashwini 2016: 19

(list).

Types: F holotype.

Habitat: “India?”

Deposited: IRSN.

Arthrosphaera marmorata (Butler, 1882)

Zephronia marmorata Butler, 1882: 197 (first description)

Zephronia marmorata :— Preudhomme de Borre: 25 (list)

Arthrosphaera marmorata :— Pocock 1899: 274 (new com¬

bination)

Arthrosphaera marmorata :— Attems 1914: 148 (list)

Arthrosphaera marmorata :— Attems 1936: 166 (list)

Arthrosphaera marmorata :— Rajulu 1970: 126 (list)

©ZFMK

Species Catalogue Sphaerotheriida

Arthrosphaera marmorata — Jeekel 2001: 10 (list)

Arthrosphaera marmorata.— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera marmorata:— Sridhar 2015: 3 (list)

Arthrosphaera marmorata:— Sridhar &Ashwini 2016: 19

(list).

Types: Syntypes (not well-preserved).

Habitat: India.

Deposited: NHML.

Arthrosphaera nitida Pocock, 1895

Arthrosphaera nitida Pocock, 1895: 411 (first description)

Arthrosphaera nitida:— Pocock 1899: 281 (list)

Arthrosphaera nitida:— Attems 1914: 148 (list)

Arthrosphaera nitida :-- Attems 1936: 166 (list)

Arthrosphaera nitida :-- Rajulu 1970: 126 (list)

Arthrosphaera nitida — Demange 1977: 231 (new local.)

Arthrosphaera nitida — Bano 1983: 168 (list)

Arthrosphaera nitida :-- Janardanan & Ramachandran

1983: 408 (parasites)

Arthrosphaera nitida — Achar 1986: 90 (cytology)

Arthrosphaera nitida — Jeekel 2001: 10 (list)

Arthrosphaera nitida :-- Sridhar & Kadamannaya 2009: 69

(list)

Arthrosphaera nitida — Sridhar 2015: 4 (list)

Arthrosphaera nitida :-- Sridhar & Ashwini 2016: 19 (list).

Types: Holotype of unknown sex.

Habitat: India: Kodaikanal, Palnai Hills.

Deposited: NHML.

Arthrosphaera noticeps (Butler, 1872)

Zephronia noticeps Butler, 1872: 355 (first description)

Zephronia noticeps — Butler 1873: 178 (description)

Zephronia noticeps — Preudhomme de Borre: 25 (list)

Arthrosphaera noticeps — Pocock 1899: 271 (new com¬

bination)

Arthrosphaera noticeps :— Attems 1914: 148 (list)

Arthrosphaera noticeps :— Attems 1936: 166 (list)

Arthrosphaera noticeps — Jeekel 2001: 10 (list)

Arthrosphaera noticeps — Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera noticeps — Sridhar 2015: 4 (list)

Arthrosphaera noticeps :— Sridhar & Ashwini 2016: 19

(list)

Arthrosphaera noticeps — De Zoysa et al. 2016: 455 (list).

Types: F holotype.

Habitat: Sri Lanka.

Deposited: NHML.

Arthrosphaera pelloceps (Chamberlin, 1921)

Trochosomapelloceps Chamberlin, 1921: 56 (first descrip¬

tion)

Trochosomapelloceps — Attems 1936: 192 (list)

Arthrosphaera pelloceps :— Jeekel 2001: 10 (new combi¬

nation).

Types: M holotype and F paratype.

Habitat: India: Periakulum, South India.

Deposited: Holotype MCZ 4694, paratype MCZ 4695.

Arthrosphaera pilifera (Butler, 1872)

Zephronia pilifera Butler, 1872: 357 (first description)

Zephronia pilifera:— Butler 1873: 180 (description)

Zephronia pilifera — Preudhomme de Borre 1884: 25 (list)

Arthrosphaera pilifera :— Pocock 1899: 270 (new comb.)

Arthrosphaera pilifera :— Attems 1914: 149 (list)

Arthrosphaera pilifera:— Attems 1936: 166 (list)

Arthrosphaera pilifera:— Jeekel 2001: 11 (list)

Arthrosphaera pilifera :— Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera pilifera — Sridhar 2015: 4 (list)

Arthrosphaera pilifera :— Sridhar & Ashwini 2016: 19

(list)

Arthrosphaera pilifera — De Zoysa et al. 2016: 455 (list).

Types: F holotype.

Habitat: Sri Lanka.

Deposited: NHML?

Arthrosphaerapygostolis Attems, 1935

Arthrosphaera pygostolis Attems, 1935: 141 (first descrip¬

tion)

Arthrosphaera pygostolis.— Attems 1936: 167 (list)

Arthrosphaera pygostolis :— Rajulu 1970: 127 (list)

Arthrosphaera pygostolis :— Jeekel 2001: 11 (list)

Arthrosphaera pygostolis — Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera pygostolis :— Sridhar 2015: 4 (list)

Arthrosphaera pygostolis :— Sridhar & Ashwini 2016: 19

(list).

Types: F holotype.

Habitat: India: Ramandroog, Madras Presidency.

Deposited: NHMW?

Arthrosphaera ruginosa Jeekel, 2001

Arthrosphaera ruginosa Jeekel, 2001: 11 (first description)

— Zephronia rugulosa :— Butler, 1872: 355 (first descrip¬

tion, e. p.)

Zephronia rugulosa :— Preudhomme de Borre 1884: 25

(list)

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

NOT Zephronia rugulosa Gervais, 1847: 82 (=

Sphaerotherium rugulosum Brandt, 1841: 363).

Arthrosphaera rugulosa :— Pocock 1899: 273 (new com¬

bination)

Arthrosphaera rugulosa:— Attems 1914: 149 (list)

Arthrosphaera rugulosa:— Attems 1936: 166 (list)

Arthrosphaera rugulosa :— Jeekel 2001: 11 (list)

Arthrosphaera rugulosa — Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera rugulosa :— Sridhar 2015: 4 (list)

Arthrosphaera rugulosa :-- Sridhar & Ashwini 2016: 19

(list)

Arthrosphaera rugulosa :— De Zoysa et al. 2016: 456 (list)

— Zephronia brandtii Pocock 1892: 143 (misidentifica-

tion), NOT Sphaeropoeus brandtii Humbert, 1865.

Remarks: According to Pocock (1899), most likely a syn¬

onym of A. brandtii.

Types: Immature holotype of unknown sex.

Habitat: Sri Lanka

Deposited: NHML.

Arthrosphaera rugosa Verhoeff) 1930

Arthrosphaera rugosa Verhoeff, 1930: 209 (first descrip¬

tion)

Arthrosphaera rugosa :— Attems 1936: 166 (list)

Arthrosphaera rugosa :— Jeekel 2001: 11 (list)

Arthrosphaera rugosa :— Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera rugosa :— Sridhar 2015: 4 (list)

Arthrosphaera rugosa :— Sridhar & Ashwini 2016: 19 (list)

Arthrosphaera rugosa :— De Zoysa et al. 2016: 456 (list).

Types: M holotype.

Habitat: Sri Lanka.

Deposited: Unknown.

Arthrosphaera scholastica Attems, 1936

Arthrosphaera scholastica Attems, 1936: 157 (first de¬

scription)

Arthrosphaera schlostica [sic!]:— Rajulu 1970: 124 (list)

Arthrosphaera scholastica— Jeekel 2001: 11 (list)

Arthrosphaera scholastica :— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera scholastica :— Sridhar 2015: 4 (list)

Arthrosphaera scholastica :— Sridhar & Ashwini 2016: 19

(list).

Types: M and F syntypes

Habitat: India: Marian Shola, Palni Hills, South India.

Deposited: NHMW 2211 (Stagl & Schiller, in prep.).

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera severa Attems, 1935

Arthrosphaera severa Attems, 1935: 139 (first description)

Arthrosphaera severa:— Attems 1936: 167 (list)

Arthrosphaera severa:— Rajulu 1970: 126 (list)

Arthrosphaera severa — Jeekel 2001: 11 (list)

Arthrosphaera severa — Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera severa:— Sridhar 2015: 4 (list)

Arthrosphaera severa:— Sridhar & Ashwini 2016: 19 (list).

Types: M and F syntypes.

Habitat: India: Ramandroog, Madras Presidency.

Deposited: NHMW 2212 (Stagl & Schiller, in prep.).

Arthrosphaera silvestrii Jeekel, 2001

— Arthrosphaera corrugata Silvestri, 1897b: 359 (first

descrip., e. p.), NOT Zephronia corrugata Butler, 1872

Arthrosphaera corrugata :— Pocock 1899: 285 (list)

Arthrosphaera corrugata:- Attems 1914: 147 (list)

Arthrosphaera corrugata :— Attems 1936: 165 (list)

Arthrosphaera corrugata :— Sridhar 2015: 3 (list).

Types: M holotype?

Habitat: “India?”.

Deposited: IRSN.

Arthrosphaera stridula (Verhoeff, 1937)

Trivandrobelum stridulum Verhoeff, 1937: 83 (first de¬

scription)

Arthrosphaera stridula :— Jeekel 2001: 11 (new comb.)

Arthrosphaera indica — Jeekel 2001: 9 (nomen nudum).

Remarks: Jeekel (2001) in his list of SE Asian

Sphaerotheriida apparently mistook an erroneous figure

label in Verhoeffs description of Trivandrobelum stridu¬

lum (Verhoeff 1937: fig. 2) stating the name

“Arthrosphaera indica” as a new description. Because the

same figure label says earlier Trivandrobelum stridulum

n. g. n. sp. and is located in the text description of T. stridu¬

lum I regard the label “Arthrosphaera indica” as a print¬

er’s error and not a valid species name.

Types: M holotype.

Habitat: India: Nedumangad, E. of Trivandrum.

Deposited: Unknown.

Arthrosphaera thurstoni Pocock, 1895

Arthrosphaera thurstoni Pocock, 1895: 411 (first descrip.)

Arthrosphaera thurstoni :— Pocock 1899: 276 (redescrip¬

tion, new locality)

Arthrosphaera thurstoni:— Attems 1914: 149 (list)

Arthrosphaera thurstoni :— Attems 1936: 166 (list)

©ZFMK

Species Catalogue Sphaerotheriida

Arthrosphaera thurstoni — Rajulu 1970: 126 (list)

Arthrosphaerathurstoni — Jeekel 2001: 11 (list)

Arthrosphaera thurstoni -- Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera thurstoni.— Sridhar 2015: 4 (list)

Arthrosphaera thurstoni.— Sridhar & Ashwini 2016: 19

(list).

Types: M and F syntypes.

Habitat: India: Nilghiri Hills; Dimhutti, near Kotagiri, Nil-

ghiri Hills; Yercand, Sheveroy Hills.

Deposited: NHML?

Arthrosphaera transitiva Attems, 1936

Arthrosphaera transitiva Attems, 1936: 160 (first descrip.)

Arthrosphaera transitiva — Rajulu 1970: 124 (list)

Arthrosphaera transitiva — Varadaraj et al. 1983: 197

(haemocytes)

Arthrosphaera transitiva — Jeekel 2001: 12 (list)

Arthrosphaera transitiva:— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera transitiva — Sridhar 2015: 4 (list)

Arthrosphaera transitiva :— Sridhar & Ashwini 2016: 19

(list).

Types: F holotype.

Habitat: India: Ganjam, Madras Presidency.

Deposited: Indian Museum Calcutta? (now Zoological

Survey of India, Kolkata).

Arthrosphaera versicolor (White, 1895)

Zephronia versicolor White, 1859: 405 (first description)

Zephronia versicolor — Humbert 1865: 41 (description)

Zephronia versicolor.— Preudhomme de Borre 1884: 25

(list)

Arthrosphaera versicolor— Pocock 1899: 272 (new com¬

bination, new locality)

Arthrosphaera versicolor— Attems 1914: 149 (list)

Arthrosphaera versicolor— Attems 1936: 167 (list)

Arthrosphaera versicolor— Jeekel 2001: 12 (list)

Arthrosphaera versicolor— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera versicolor— Sridhar 2015: 4 (list)

Arthrosphaera versicolor— Ambarish & Sridhar 2015c:

(distribution)

Arthrosphaera versicolor— Sridhar & Ashwini 2016: 19

(list)

Arthrosphaera versicolor — De Zoysa et al. 2016: 456

(list).

Types: M holotype?

Habitat: Sri Lanka: Peradenia.

Deposited: NHML?

Bonn zoological Bulletin - Supplementum 63: 1-104

Arthrosphaera wroughtoni Pocock, 1895

Arthrosphaera wroughtoni Pocock, 1895: 411 (first de¬

scription)

Arthrosphaera wroughtoni — Pocock 1899: 275 (descrip¬

tion)

Arthrosphaera wroughtoni :— Attems 1914: 149 (list)

Arthrosphaera wroughtoni :— Attems 1936: 167 (list)

Arthrosphaera wroughtoni :— Rajulu 1970: 126 (list)

Arthrosphaera wroughtoni — Jeekel 2001: 12 (list)

Arthrosphaera wroughtoni :— Sridhar & Kadamannaya

2009: 69 (list)

Arthrosphaera wroughtoni — Sridhar 2015: 4 (list)

Arthrosphaera wroughtoni — Sridhar & Ashwini 2016: 19

(list).

Types: M holotype.

Habitat: India: Kanara.

Deposited: NHML?

Arthrosphaera zebraica (Butler, 1872)

Zephronia zebraica Butler, 1872: 356 (first description)

Zephronia zebraica:— Preudhomme de Borre 1884: 25

(list)

Arthrosphaera zebraica — Pocock, 1899: 275 (new com¬

bination)

Arthrosphaera zebraica - Attems 1914: 149 (list)

Arthrosphaera zebraica - Attems 1936: 167 (list)

Arthrosphaera zebriaca [sic!]:— Karandikar & Rodgi

1955: 27 (parasites)

Arthrosphaera zebraica:— Chowdaiah 1966: 847 (cytol¬

ogy)

Arthrosphaera zebraica:— Rajulu 1970: 126 (list)

Arthrosphaera zebraica:— Kunte 1974: 673 (list)

Arthrosphaera zebraica :— Rangaswamy et al. 1978: 98

(distribution)

Arthrosphaera zebraica:— Bano 1983: 168 (list)

Arthrosphaera zebraica :— Rangaswamy et al. 1983: 7

(distribution)

Arthrosphaera zebraica — Jeekel 2001: 12 (list)

Arthrosphaera zebraica :— Sridhar & Kadamannaya 2009:

69 (list)

Arthrosphaera zebraica — Kadamannaya et al. 2012: 183

(morphology)

Arthrosphaera zebraica:— Ambarish & Sridhar 2013a: 64

(distribution, behaviour)

Arthrosphaera zebraica — Sridhar 2015: 4 (list)

Arthrosphaera zebraica :— Sridhar & Ashwini 2016: 19

(list).

Types: M holotype.

Habitat: India: Bombay.

Deposited: NHML.

©ZFMK

Thomas Wesener

Genus Sphaeromimus

de Saussure & Zehntner. 1902

Sphaeromimus de Saussure & Zehntner, 1902: 20 (first

description); Attems 1926: 119 (list); Attems 1942: 60

(list); Jeekel 1971: 28 (list); Jeekel 1974: 45 (classifica¬

tion); Hoffman 1980: 63 (list); Jeekel 1999: 8 (catalogue,

discussion); Enghoff 2003: 618 (list); Wesener & Sierwald

2005b: 557 (redescription, additional species); Wesener &

Wagele 2007: 147 (ecology); Wesener 2009: 8 (key); We¬

sener & VandenSpiegel 2009: 548 (morphological phylo¬

genetic analysis); Wesener et al. 2010b: 1185 (molecular

phylogenetic analysis); Wesener 2014a: 199 (morpholo¬

gical phylogenetic analysis); Wesener et al. 2014: 72 (re¬

vision, additional species).

Description: Anterior telopod with four podomeres dis¬

tal to syncoxite, telopoditomere 4 always with three spines

in special positions. Harp carrying 3-10 stridulation ribs.

Posterior telopods with three telopoditomeres, 3rd carry¬

ing two membranous lobes. Operculum of vulva greatly

enlarged and well-rounded, often larger than bursa. Inner

plate of vulva not protruding. Female washboard with

8-20 stridulation ribs. Legs with a coxal lobe, tarsi very

wide, paddle-like. Underside of anal shield with a single,

short, locking carina. First antennomere with neither a

groove nor scales, nor sensilla basiconica. Antennomere

6 large and flattened, but not axe-shaped. Number of api¬

cal cones sexually dimorphic, females with 40-50, males

with 80-90. 20-40 mm long.

Type species: Sphaeropoeus musicus de Saussure &

Zehntner, 1897, by monotypy.

Distribution: S and SE Madagascar

Species included (10)

Sphaeromimus andohahela Wesener, 2014

Sphaeromimus andrahomana Wesener, 2014

Sphaeromimus inexpectatus Wesener & Sierwald, 2005

Sphaeromimus ivohibe Wesener, 2014

Sphaeromimus lavasoa Wesener, 2014

Sphaeromimus musicus (de Saussure & Zehntner, 1897)

Sphaeromimus saintelucei Wesener, 2014

Sphaeromimus splendidus Wesener & Sierwald, 2005

Sphaeromimus titanus Wesener, 2014

Sphaeromimus vatovavy Wesener, 2014

Sphaeromimus andohahela Wesener, 2014

(Fig. 13F)

Sphaeromimus ‘n. sp. I Manantantely’; Sphaeromimus ‘n.

sp. II Malio’:-- Wesener et al. 2010: 1185 (molecular

phylogenetic analysis)

Sphaeromimus andohahela Wesener et al. 2014: 89 (first

description).

Bonn zoological Bulletin - Supplementum 63: 1-104

Types: M holotype, 4 F, 3 imm. paratypes.

Localities: Madagascar, Province Toliara, PN Andohahela,

Isaka-Ivondro Nord (Type), Malio, Manantantely

Deposited: ZFMK MYR2322 (holotype); FMNH-INS

61135 (1 F paratype); FMNH-INS 61136 (1 imm. para-

type); FMNH-INS 61137 (1 F paratype); FMN-INS 56212

(2 F, 2 imm. paratypes)

Sphaeromimus andrahomana Wesener, 2014

Sphaeromimus ‘n. sp. V Grotte’:-- Wesener et al. 2010:

1185 (molecular phylogenetic analysis)

Sphaeromimus andrahomana Wesener et al. 2014: 98 (first

description).

Types: M holotype.

Localities: Madagascar, Province Toliara, N of Ankapaky

(type), Grotte d’Andrahomana.

Deposited: FMNH-INS 56214 (holotype).

Sphaeromimus inexpectatus Wesener & Sierwald, 2005

(Fig. 13D)

Sphaeromimus inexpectatus Wesener & Sierwald, 2005b:

570 (first description)

Sphaeromimus inexpectatus.— Wesener & Wagele 2007:

150 (ecology)

Sphaeromimus inexpectatus — Wesener 2009: 131 (list)

Sphaeromimus inexpectatus :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaeromimus inexpectatus :— Wesener et al. 2010b: 1185

(molecular phylogenetic analysis)

Sphaeromimus inexpectatus :— Temple et al. 2012: 37:

1185 (conservation)

Sphaeromimus inexpectatus :— Wesener 2014a: 199 (mor¬

phological phylogenetic analysis)

Sphaeromimus inexpectatus :— Wesener et al. 2014: 76 (ad¬

ditional specimens).

Types: M holotype, 1 F paratype (fragmented).

Localities: Madagascar, Province Toliara, Mandena littoral

rainforest (Type); Enato.

Deposited: FMNH-INS 6701.

Sphaeromimus ivohibe Wesener, 2014

Sphaeromimus ivohibe Wesener et al. 2014: 93 (first de¬

scription).

Types: M holotype.

Localities: Madagascar, Province Fianarantsoa, RS de Ivo¬

hibe.

Deposited: FMNH-INS 8184.

©ZFMK

Species Catalogue Sphaerotheriida

Sphaeromimus lavasoa Wesener, 2014

(Fig. 13E)

Sphaeromimus ‘n. sp. Ill G-Lavasoa’:-- Wesener et al.

2010: 1185 (molecular phylogenetic analysis)

Sphaeromimus lavasoa Wesener et al. 2014: 84 (first de¬

scription).

Types: M holotype, 40 M & F paratypes.

Localities: Madagascar, Province Toliara, Ambitsoron-

gorongo, Grande Lavasoa (Type), Petit Lavasoa.

Deposited: ZFMK MYR2320 (holotype); ZFMK

MYR2321 (2 F paratypes); FMNH-INS 61141 (1 M

paratype); FMNH-INS 61134 (1 M paratype); FMNH-INS

61142 (1 F paratype); FMN-INS 61143 (1 F paratype);

FMNH-INS 56213 (1 F paratype); FMNH-INS 56208 (35

M & F paratypes).

Sphaeromimus musicus (de Saussure & Zehntner, 1897)

(Figs 12B, 13 A)

Sphaeropoeus musicus de Saussure & Zehntner, 1897: pi.

4, fig. 1 a-e (first description)

Sphaeromimus musicus.— de Saussure & Zehntner 1902:

75 (description)

Sphaeromimus musicus :— Jeekel 1999: 8 (list)

Sphaeromimus musicus :— Enghoff 2003: 618 (list)

Sphaeromimus musicus :— Wesener & Sierwald 2005b:

564 (redescription)

Sphaeromimus musicus — Wesener 2009: 131 (list)

Sphaeromimus musicus :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaeromimus musicus :— Wesener et al. 2010b: 1185

(molecular phylogenetic analysis)

Sphaeromimus musicus — Wesener 2014a: 199 (morpho¬

logical phylogenetic analysis)

Sphaeromimus musicus :— Wesener et al. 2014: 75 (revi¬

sion, new localities).

Types: M holotype.

Localities: Madagascar (Type), S Madagascar spiny for¬

est ecosystem.

Deposited: MNHN CH038.

Sphaeromimus saintelucei Wesener, 2014

Sphaeromimus saintelucei Wesener et al. 2014: 96 (first

description).

Types: M holotype, 2 M, 1 ? paratypes.

Localities: Madagascar, Province Toliara, Sainte Luce lit¬

toral rainforest, fragment S9.

Deposited: ZFMK MYR889 (holotype); FMNH-INS

61089 (1 M paratype); FMNH-INS 61088 (1 ? paratype).

Sphaeromimus splendidus Wesener & Sierwald, 2005

(Fig. 13C)

Sphaeromimus splendidus Wesener & Sierwald, 2005b:

567 (first description)

Sphaeromimus splendidus — Wesener & Wagele 2007:

150 (ecology)

Sphaeromimus splendidus — Wesener 2009: 131 (list)

Sphaeromimus splendidus :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Sphaeromimus splendidus :— Wesener et al. 2010b: 1185

(molecular phylogenetic analysis)

Sphaeromimus splendidus :— Wesener 2014a: 199 (mor¬

phological phylogenetic analysis)

Sphaeromimus splendidus :— Wesener et al. 2014: 75 (ad¬

ditional specimens).

Types: 1 F holotype, 1 M, 3 F paratypes.

Localities: Madagascar, Province Toliara, Sainte Luce lit¬

toral rainforest, fragment S9.

Deposited: FMNH-INS 6702 (holotype); 1 M, 2 F

paratypes FMNH-INS 6703; 1 F paratype CAS.

Sphaeromimus titanus Wesener, 2014

Sphaeromimus titanus Wesener et al. 2014: 77 (first de¬

scription).

Types: M holotype, F paratype.

Localities: Madagascar, Province Fianarantsoa, Manom-

bo.

Deposited: CAS ENT 9032789.

Sphaeromimus vatovavy Wesener, 2014

Sphaeromimus vatovavy Wesener et al. 2014: 81 (first de-

scrition).

Types: M holotype, F paratype.

Localities: Madagascar, Province Fianarantsoa, Tsianovo-

ha (Fort Camot).

Deposited: MNHN ‘39’.

Genus Microsphaerotherium

Wesener & VandenSpiegel 2007

Microsphaerotherium Wesener & VandenSpiegel, 2007:

153 (first description); Wesener 2009: 8 (key); Wesener

& VandenSpiegel 2009: 548 (morphological phylogenet¬

ic analysis); Wesener 2014a: 199 (morphological phylo¬

genetic analysis).

Description: Anterior telopod with three podomeres dis¬

tal to syncoxite. Harp carrying a single stridulation rib.

Posterior telopods with three telopoditomeres, 3rd with a

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

single membranous lobe. Operculum of vulva narrow, in¬

ner plate of vulva protruding. 4-6 discontinuous ribs on

female washboard. Legs without a coxal lobe, tarsi always

long and slender. Underside of anal shield with a single

(often fused) locking carina, this being very long, some¬

times connecting with the locking carina from the other

side. Surface of tergites covered with special tiny ‘rotat¬

ed’ (helicoid) setae, dorsally with a sclerotized crest. First

antennomere with a groove, few sensilla basiconica, and

isolated sclerotized scales. All antennomeres cylindrical,

with 5-10 apical cones. Smallest known sphaerotheriids.

Type species: Microsphaerotherium ivohibiense Wesen¬

er & VandenSpiegel, 2007, by monotypy.

Distribution: Central Madagascar

Species included (2)

Microsphaerotherium anjozorobe Wesener 2009

Microsphaerotherium ivohibiense Wesener & VandenSpie¬

gel, 2007

Microsphaerotherium anjozorobe Wesener, 2009

Microsphaerotherium anjozorobe Wesener, 2009: 9 (first

description).

Types: M holotype, 1 M paratype.

Localities: Madagascar, Province d’Antananarivo, 3 km

NE Andranomay, 1300 m.

Deposited: CAS BLF 2370 & 2371.

Microsphaerotherium ivohibiense Wesener & Vanden¬

Spiegel, 2007

Microsphaerotherium ivohibiensis [sic!] Wesener & Van¬

denSpiegel, 2007: 154 (first description)

Microsphaerotherium ivohibiense.— Wesener 2009: 131

(list)

Microsphaerotherium ivohibiense :— Wesener & Vanden¬

Spiegel 2009: 548 (morpholog. phylogenetic analysis)

Microsphaerotherium ivohibiense.— Wesener 2014a: 199

(morphological phylogenetic analysis).

Types: M holotype, 1 M paratype.

Localities: Madagascar, Province Fianarantsoa, RS Ivo-

hibe, 1200 m.

Deposited: FMNH-INS 4011.

Genus Zoosphaerium Pocock. 1895

(Figs 14A-H)

Zoosphaerium Pocock, 1895: 410 (first description).

Chamberlin 1921: 59 (additional species); Jeekel 1971: 29

(list); Jeekel 1974: 45 (classification); Hoffman 1980: 63

(list); Jeekel 1999: 5 (catalogue); Thorstrom & Lind 1999:

572 (predation birds); Shelley et al. 2000: 13 (catalogue);

Enghoff 2003: 618 (list); Wesener & Sierwald 2005a: 3

(additional species); Wesener & Wagele 2007: 147 (ecol¬

ogy); Wesener & Wagele 2008: 9 (redescription); Wesen¬

er 2009: 1 (additional species); Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis); Wesen¬

er et al. 2010a: 133 (additional species); Wesener et al.

2010b: 1185 (molecular phylogenetic analysis); Wesener

2014a: 199 (morphological phylogenetic analysis); Koch

2015: 12 (morphology).

Synonyms:

— Globotherium Brolemann, 1922: 230 (first descrip¬

tion), tentatively synonymized with Zoosphaerium by

Jeekel 1999: 9. Attems 1926: 119 (list); Attems 1942:

60 (list); Jeekel 1971: 29 (list); Jeekel 1974: 45 (clas¬

sification); Hoffman 1980: 63 (list); Enghoff 2003: 619,

tables 8-19 (all Globotherium species assigned to

Zoosphaerium ); Wesener & Wagele 2008: 9 (formal

synonymization with Zoosphaerium ).

Type species : Sphaerotherium digitale de Saussure &

Zehntner, 1897, synonymized under Sphaerotherium

neptunus Butler, 1872 by Wesener & Wagele 2008: 10.

— Heligmasoma Chamberlin, 1921: 58 (first description),

formal synonymization with Zoosphaerium by Wesen¬

er & Wagele 2008: 9. Jeekel 1971: 29 (list); Jeekel 1974:

45 (classification); Hoffman 1980: 63 (list).

Type species : Heligmasoma errans Chamberlin, 1921,

synonymized under Sphaerotherium coquerelianum de

Saussure & Zehntner, 1897 by Wesener & Wagele 2008:

38.

Description: Anterior telopod with three podomeres dis¬

tal to syncoxite. Harp carrying one or two stridulation ribs

(one species with three). Posterior telopods with three

telopoditomeres, 3rd extremely variable, but at least in one

species group (the libidinosum group) carrying two mem¬

branous lobes. Operculum of vulva of variable size, but

apical margin always with a median notch. Inner plate of

vulva protruding. Female washboard with 2-8 stridulation

ribs. Legs without a coxal lobe, tarsi always long and slen¬

der. Underside of anal shield with two locking carinae,

posterior one usually 2-3 times larger than anterior one.

First antennomere with a groove, covered with sclerotized

scales. Isolated sensilla basiconica are always present on

first antennomere, rarely also on antennomere 4 or 5. All

antennomeres cylindrical, with four or 15-50 apical cones.

Usually 20-50, giant species 70-100 mm long.

Type species: Sphaerotherium neptunus Butler, 1872, by

subsequent designation by Pocock 1895.

Distribution: Madagascar

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Species Catalogue Sphaerotheriida

Species included (54 & 11 nomina dubia)

Zoosphaerium campanulatum (de Saussure & Zehntner,

1897)

Zoosphaerium neptunus (Butler, 1872)

Zoosphaerium album Wesener, 2009

Zoosphaerium anomalum (de Saussure & Zehntner,

1902)

Zoosphaerium arborealis Wesener & Sierwald, 2005

Zoosphaerium anreum Wesener, 2009

Zoosphaerium discolor Wesener, 2009

Zoosphaerium endemicum Wesener, 2009

Zoosphaerium haackeri Wesener, 2009

Zoosphaerium lambertoni (Brolemann, 1922)

Zoosphaerium libidinosum (de Saussure & Zehntner,

1897)

Zoosphaerium mitoho Wesener, 2009

Zoosphaerium priapus (de Saussure & Zehntner, 1897)

Zoosphaerium pseudopriapus Wesener, 2009

Zoosphaerium pulchellum Wesener, 2009

Zoosphaerium xerophilum Wesener, 2009

Zoosphaerium broelemanni Wesener, 2009

Zoosphaerium corystoides Wesener, 2009

Zoosphaeriumplatylabum (de Saussure & Zehntner, 1897)

Zoosphaerium pseudoplatylabum Wesener, 2009

Zoosphaerium solitarium Wesener, 2009

Zoosphaerium tsingy Wesener, 2009

Zoosphaerium alluaudi (de Saussure & Zehntner, 1902)

Zoosphaerium amabile Wesener, 2009

Zoosphaerium ambrense Wesener, 2009

Zoosphaerium bambusoides Wesener & Bespalova, 2010

Zoosphaerium bilobum Wesener, 2009

Zoosphaerium coquerelianum (de Saussure & Zehntner,

1897)

Zoosphaerium denticulatum Wesener, 2009

Zoosphaerium ignotum Wesener, 2009

Zoosphaerium isalo Wesener, 2009

Zoosphaerium muscorum Wesener & Bespalova, 2010

Zoosphaerium smaragdinum Wesener, 2009

Zoosphaerium tainkintana Wesener, 2009

Zoosphaerium tampolo Wesener, 2009

Zoosphaerium tigrioculatum Wesener & Bespalova, 2010

Zoosphaerium trichordum Wesener, 2009

Zoosphaerium villosum Wesener & Sierwald, 2005

Zoosphaerium viridissimum Wesener, 2009

Zoosphaerium voeltzkowianum (de Saussure & Zehntner,

1897)

Zoosphaerium blandum (de Saussure & Zehntner, 1897)

Zoosphaerium pseudoblandum Wesener, 2009

Zoosphaerium micropiligerum Wesener, 2009

Zoosphaeriumpiligerum (de Saussure & Zehntner, 1897)

Zoosphaerium pseudopiligerum Wesener, 2009

Zoosphaerium analavelona Wesener, 2009

Zoosphaerium fisheri Wesener, 2009

Zoosphaerium darthvaderi Wesener & Bespalova, 2010

Zoosphaerium heleios Wesener & Bespalova, 2010

Zoosphaerium anale (de Saussure & Zehntner, 1897)

Zoosphaerium fraternarium Jeekel, 1999

Zoosphaerium immane (Karsch, 1881)

Zoosphaerium sakanum (Attems, 1910)

Zoosphaerium subreflexum Jeekel, 1999

Nomina dubia

Zoosphaerium actaeon (White, 1859)

Zoosphaerium crassum (Butler, 1878)

Zoosphaerium elegans (Lenz, 1880)

Zoosphaerium glabrum (Butler, 1873)

Zoosphaerium hippocastanum (Gervais, 1847)

Zoosphaerium latum (Butler, 1872)

Zoosphaerium lamprinum (Butler, 1878)

Zoosphaerium reflexion (Brolemann, 1922)

Zoosphaerium reticulatum (Butler, 1878)

Zoosphaerium stigmaticum (Butler, 1873)

Zoosphaerium testaceum (Olivier, 1792)

The Zoosphaerium neptunus species-group

Wesener & Wagele 2008: 9 (first description)

Zoosphaerium neptunus (Butler, 1872)

Sphaerotherium Neptunus Butler, 1872: 358, pi. 18, fig.

6 (first description)

Sphaerotherium neptunus.— Butler 1873: 176 (list)

Sphaerotherium Neptunus :— Preudhomme de Borre 1884:

27 (catalogue)

Sphaerotherium neptunus :— de Saussure & Zehntner

1902: 67 (redescription)

Zoosphaerium neptunus :— Pocock 1895: 410 (type

species of Zoosphaerium)

Zoosphaerium neptunus.— Jeekel 1999: 10 (list)

Zoosphaerium neptunus :— Enghoff 2003: 620 (list)

Zoosphaerium neptunus — Wesener & Wagele 2008: 10

(redescription)

Zoosphaerium neptunus :— Wesener 2009: 131 (list)

Zoosphaerium neptunus :— Wesener et al. 2010a: 159 (new

locality)

Zoosphaerium neptunus — Wesener & Schutte 2010: 89

(behaviour)

— Sphaerotherium digitale de Saussure & Zehntner,

1897: pi. 5, fig. 16 (first description), synonymization

under Sphaerotherium neptunus by Wesener & Wage¬

le 2008: 10

Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Thomas Wesener

Sphaerotherium digitale — de Saussure & Zehntner 1902:

54 (description)

Sphaerotherium (Globotherium) digitale.— Brolemann

1922: 232, figs 6-8 (type species of subgenus Globoth¬

erium)

Zoosphaerium digitale — Jeekel 1999: 13 (new combina¬

tion, list)

Zoosphaerium digitale — Enghoff 2003: 620 (list)

— Sphaerotherium pygidiale de Saussure & Zehntner,

1897: pi. 5, figs 8-8d (first description), synonymiza-

tion under Sphaerotherium neptunus by Wesener & Wa-

gele 2008: 10, with lectotype designation.

Sphaerotherium pygidiale :-- de Saussure & Zehntner

1902: 42, pi. 15, fig. 22 (description)

Zoosphaerium pygidiale :-- Jeekel 1999: 11 (new combi¬

nation, list)

Zoosphaerium pygidiale :— Enghoff 2003: 621 (list)

— Sphaerotherium lamprinum :— de Saussure & Zehnt¬

ner 1897: pi. 5, figs 17-17b (redescription, misidenti-

fication). NOT S. lamprinum Butler, 1878.

— Sphaerotherium latum — de Saussure & Zehntner

1897: pi. 5 , fig. 14 (redescription, misidentification).

NOT S. latum Butler, 1872.

— Sphaerotherium hippocastanum :— de Saussure &

Zehntner 1897: pi. 5, fig. 15 (redescription, misidenti¬

fication). NOT S. hippocastanum Gervais, 1847.

Types: 1 F holotype (broken).

Localities: Madagascar.

Deposited: NHML BM.1858.85.

Zoosphaerium campanulatum (de Saussure & Zehntner,

1897)

Sphaerotherium campanulatum de Saussure & Zehntner,

1897: pi. 4, figs 6-6b (first description)

Sphaerotherium campanulatum :— de Saussure & Zehnt¬

ner 1902: 28 (description)

Sphaerotherium campanulatum — Attems 1910: 75 (new

locality)

Zoosphaerium campanulatum :— Jeekel 1999: 9 (new com¬

bination)

Zoosphaerium campanulatum :— Enghoff2003: 620 (list)

Zoosphaerium campanulatum :— Wesener & Wagele

2008: 19 (redescription)

Zoosphaerium campanulatum — Wesener 2009: 131 (list).

Types: M holotype.

Localities: Madagascar, cote sud-ouest.

Deposited: MNHN CB013.

The Zoosphaerium libidinosum species-group

Wesener & Wagele 2008: 9 (key)

Zoosphaerium album Wesener, 2009

Zoosphaerium album Wesener, 2009: 18 (first descrip.).

Types: M holotype, numerous paratypes.

Localities: Madagascar, Province Toliara, Foret Vohibasia.

Deposited: FMNH-INS 5457.

Zoosphaerium anomalum (de Saussure & Zehntner,

1902)

Sphaerotherium anomalum de Saussure & Zehntner, 1902:

56 (first description)

Zoosphaerium anomalum — Jeekel 1999: 13 (new com¬

bination)

Zoosphaerium anomalum :— Enghoff 2003: 620 (list)

Zoosphaerium anomalum :— Wesener & Wagele 2008: 34

(redescription)

Zoosphaerium anomalum :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium anomalum :— Wesener 2009: 131 (list).

Types: M holotype.

Localities: Madagascar, Province Toliara, Isaka.

Deposited: MNHN CB009.

Zoosphaerium arborealis Wesener & Sierwald, 2005

Zoosphaerium arborealis Wesener & Sierwald, 2005a: 34

(first description)

Zoosphaerium arborealis :— Wesener & Wagele 2007: 148

(distribution)

Zoosphaerium arborealis :— Wesener & Wagele 2008: 28

(list)

Zoosphaerium arborealis :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium arborealis :— Wesener 2009: 131 (list).

Types: M holotype, 6 M, 17 F paratypes.

Localities: Madagascar, Tolagnaro, littoral forests of Man-

dena and Sainte Luce (type localities); PN Andringitra,

Foret d’Ravaro.

Deposited: Holotype FMNH-INS 8652. Paratypes: 2 M,

8 F FMNH-INS 56028; 1 M, 2 F CAS; 1 M, 2 F ZSM; 3

F SMNG VNR 13554; 2 M, 2 F FMNH-INS 8651.

Zoosphaerium aureum Wesener, 2009

Zoosphaerium aureum Wesener, 2009: 43 (first descrip.).

Types: M holotype, 4 M, 4 F paratypes.

Localities: Madagascar, Province Antsiranana, Montagne

d’Ambre.

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©ZFMK

Species Catalogue Sphaerotheriida

Deposited: Holotype and 3 M, 3 F paratypes ZMUC; 1

M, 1 F paratypes FMNH.

Zoosphaerium discolor Wesener, 2009

Zoosphaerium discolor Wesener, 2009: 22 (first descrip¬

tion).

Types: M holotype, numerous paratypes.

Localities: Madagascar, Province Mahajanga, PN Tsingy

de Bemaraha.

Deposited: FMNH-INS 5454.

Zoosphaerium endemicum Wesener, 2009

Zoosphaerium endemicum Wesener, 2009: 37 (first de¬

scription).

Types: M holotype.

Localities: Madagascar, Province Toliara, Foret

Analavelona.

Deposited: FMNH-INS 5420.

Zoosphaerium haackeri Wesener, 2009

Fig. 14A

Zoosphaerium haackeri Wesener, 2009: 16 (first descrip.)

Zoosphaerium haackeri -- Wesener et al. 2010b: 1187

(molecular phylogeny).

Types: M holotype.

Localities: Madagascar, Province Toliara, RS Cap Sainte

Marie.

Deposited: CAS BLF 5499.

Zoosphaerium lamberloni (Brolemann, 1922)

Sphaerotherium ( Globotherium ) lambertoni Brolemann,

1922: 235 (first description)

Zoosphaerium lambertoni.— Jeekel 1999: 13 (new com¬

bination)

Zoosphaerium lambertoni :— Enghoff 2003: 620 (list)

Zoosphaerium lambertoni :— Wesener & Wagele 2008: 28

(list)

Zoosphaerium lambertoni :— Wesener 2009: 131 (list).

Types: M holotype.

Localities: Madagascar, surroundings of Antananarivo.

Deposited: Academie Malgache (now zoological collec¬

tion Tsimbazawa), presumably lost.

Zoosphaerium libidinosum (de Saussure & Zehntner, 1897)

Sphaerotherium libidinosum de Saussure & Zehntner,

1897: pi. 4, fig. 2 (first description)

Bonn zoological Bulletin - Supplementum 63: 1-104

Sphaerotherium libidinosum.— de Saussure & Zehntner

1901: 435 (new localities)

Sphaerotherium libidinosum :— de Saussure & Zehntner

1902: 69 (description)

Sphaerotherium libidinosum :— Attems 1910: 80 (list)

Sphaerotherium ( Globotherium ) libidonosum — Demange

1969: 485 (new locality)

Zoosphaerium libidinosum :— Jeekel 1999: 12 (new com¬

bination)

Zoosphaerium libidinosum :— Enghoff 2003: 620 (list)

Zoosphaerium libidinosum.— Wesener & Wagele 2008: 22

(redescription)

Zoosphaerium libidinosum — Wesener 2009: 131 (list)

Zoosphaerium libidinosum :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis).

Types: 1 M lectotype, 5 M paralectotypes (designated by

Wesener & Wagele 2008: 22).

Localities: Madagascar (Type), widespread in SW Mada¬

gascar spiny forest zone.

Depository: MNHN CB035.

Zoosphaerium mitoho Wesener, 2009

Zoosphaerium mitoho Wesener, 2009: 27 (first descrip¬

tion).

Types: M holotype, 1 M, 1 F paratypes.

Localities: Madagascar, Province Toliara, PN Tsimanam-

petsotsa, Mitoho Cave.

Depository: Type series CAS BLF 6163.

Zoosphaerium priapus (de Saussure & Zehntner, 1897)

Sphaerotherium priapus de Saussure & Zehntner, 1897:

pi. 4, fig. 5 (first description)

Sphaerotherium priapus — de Saussure & Zehntner,

1902: 61 (description)

Zoosphaerium priapus :— Jeekel 1999: 14 (new combina¬

tion)

Zoosphaerium priapus — Enghoff 2003: 620 (list)

Zoosphaerium priapus — Wesener & Wagele 2008: 30 (re¬

description)

Zoosphaerium priapus — Wesener 2009: 131 (list)

Zoosphaerium priapus — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis).

Types: 1 F lectotype (designated by Wesener & Wagele

2008: 30).

Localities: Madagascar, Antananarivo (Type); Foret d’O-

rangea.

Depository: MNHN CB042.

©ZFMK

Thomas Wesener

Zoosphaerium pseudopriapus Wesener, 2009

Zoosphaerium pseudopriapus Wesener, 2009: 46 (first de¬

scription)

Zoosphaerium pseudopriapus.— Wesener et al. 2010a: 159

(new locality).

Types: M holotype, 3 M, 4 F paratypes.

Localities: Madagascar, Province Antsiranana, Foret de

Bekaraoka (Type); Analamozava River near Daraina;

Foret d’Ampondrabe; Foret de Binara.

Depository: CAS BLF 9871.

Zoosphaerium pulchellum Wesener, 2009

Zoosphaerium pulchellum Wesener, 2009: 33 (first de¬

scription).

Types: M holotype, 1 M, 1 F paratypes.

Localities: Madagascar, Province Toliara, Andohahela par¬

cel 1, 15 km NW Eminiminy.

Depository: FMNH-INS 5388.

Zoosphaerium xerophilum Wesener, 2009

Zoosphaerium xerophilum Wesener, 2009: 40 (first de¬

scription)

Zoosphaerium xerophilum :— Wesener et al. 2010a: 157

(description of female).

Types: M holotype, 2 M paratypes.

Localities: Madagascar, Province Antsiranana, RS d’Am-

bre.

Depository: FMNH-INS 3935.

The Zoosphaerium platvlabum species-group

Wesener & Wagele 2008: 50

Zoosphaerium hroelemanni Wesener, 2009

Zoosphaerium hroelemanni Wesener, 2009: 70 (first des¬

cription)

Zoosphaerium hroelemanni — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis).

— Sphaerotherium (Globotherium) hippocastanum :—

Brolemann 1922: 230 (description, misidentification),

NOT Sphaerotherium hippocastanum Gervais, 1847.

Types: M holotype, 3 M, 3 F paratypes.

Localities: Madagascar, Province Antananarivo, Antana¬

narivo.

Depository: MNHN CB029, holotype, 2 M, 3 F paratypes;

MNHN CB030, 1 M paratype.

Zoosphaerium corystoides Wesener, 2009

Zoosphaerium corystoides Wesener, 2009: 66 (first de¬

scription) .

Zoosphaerium corystoides - Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis).

Types: M holotype, 30 M, 4 F paratypes.

Localities: Madagascar, Province Antsiranana, Montagne

d’Ambre.

Depository: FMNH-INS 3932, holotype, 27 M, 3 F

paratypes; CAS BLF 2782, 3 M, 1 F paratypes.

Zoosphaerium platylabum (de Saussure & Zehntner,

1897)

Sphaerotherium platylabum de Saussure & Zehntner,

1897: pi. 5, fig. 9 (first description)

Sphaerotherium platylabum — de Saussure & Zehntner

1902: 54 (description)

Sphaerotherium platylabum — Attems 1910: 86 (new lo¬

cality)

Sphaerotherium ( Globotherium ) platylabum :— Brole¬

mann 1922: 230 (list)

Zoosphaerium platylabum — Jeekel 1999: 13 (new com¬

bination)

Zoosphaerium platylabum — Enghoff 2003: 620 (list)

Zoosphaerium platylabum :— Wesener & Wagele 2008: 50

(redescription)

Zoosphaerium platylabum — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium platylabum — Wesener et al. 2010a: 159

(additional specimens).

Types: M holotype.

Localities: Madagascar, Province Antananarivo, Antana¬

narivo (Type); Andasibe, Perinet.

Depository: MNHN CB041.

Zoosphaerium pseudoplatylabum Wesener, 2009

Zoosphaerium pseudoplatylabum Wesener, 2009: 54

(first description)

Zoosphaerium pseudoplatylabum — Wesener et al. 2010a:

158 (description of female).

Types: M holotype, 3 M paratypes.

Localities: Madagascar, Province Fianarantsoa, Foret de

Vinantelo (type); Andringitra; RS de Ivohibe.

Depository: FMNH-INS 5425, holotype & 1 paratype;

FMNH-INS 5421, 2 paratypes.

Zoosphaerium solitarium Wesener, 2009

Zoosphaerium solitarium Wesener, 2009: 58 (first descrip¬

tion).

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©ZFMK

Species Catalogue Sphaerotheriida

Types: M holotype, 3 M, 1 F paratypes.

Localities: Madagascar, Province Antsiranana, Montagne

des Fran^ais.

Depository: ZSM FGZC1897, holotype; Paratypes:

ZMUH FGZC1662; ZSM FGZC1842; ZSM FGZC1693

(1 F); CAS BLF3129.

Zoosphaerium tsingy Wesener, 2009

Zoosphaerium tsingy Wesener, 2009:61 (first description).

Types: M holotype, 13 M, 7 F paratypes.

Localities: Madagascar, Province Antsiranana, Tsingy de

Bemaraha.

Depository: Holotype, 8 M, 2 F paratypes, FMNH-INS

5455; 5 M, 5 F paratypes, FMNH-INS 5454.

The Zoosphaerium coquerelianum

species-group

synonym Zoosphaerium alluaudi species-group

Wesener & Sierwald 2005a: 8.

Zoosphaerium alluaudi (de Saussure & Zehntner, 1902)

(Fig. 14G)

Sphaerotherium alluaudi de Saussure & Zehntner, 1902:

58 (first description)

Zoosphaerium alluaudi.— Jeekel 1999: 13 (new comb.)

Zoosphaerium alluaudi :— Enghoff 2003: 620 (list)

Zoosphaerium alluaudi — Wesener & Sierwald 2005a: 8

(redescription)

Zoosphaerium alluaudi — Wesener & Wagele 2007: 148

(distribution)

Zoosphaerium alluaudi :— Wesener & Wagele 2008:42 (list)

Zoosphaerium alluaudi :— Wesener 2009: 131 (list)

Zoosphaerium alluaudi :— Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium alluaudi:- Wesener et al. 2010b: 1187 (mo¬

lecular phylogenetic analysis)

Zoosphaerium alluaudi :— Temple et al. 2012: 37: 1185

(conservation).

Types: M lectotype, 2 F paralectotypes (designated We¬

sener & Sierwald 2005a: 8).

Localities: Madagascar, Fort Dauphin (Type); Petriky and

Lavasoa Mountains.

Depositories: Lectotype and paralectotypes MNHN

CB006.

Zoosphaerium amabile Wesener, 2009

Zoosphaerium amabile Wesener, 2009: 74 (first descrip.)

Zoosphaerium amabile :— Wesener et al. 2010a: 158 (ad¬

ditional specimens).

Types: M holotype.

Localities: Madagascar, Province Toamasina, Montagne

d’Akirindro.

Depositories: Holotype CAS BLF 8254.

Zoosphaerium ambrense Wesener, 2009

Zoosphaerium ambrense Wesener, 2009: 84 (first descrip.)

Zoosphaerium ambrense :— Wesener et al. 2010a: 158

(new locality).

Types: M holotype, 1 M, 2 F paratypes.

Localities: Madagascar, PN Montagne d’Ambre (Type).

Depositories: Type series, MHNG ‘Mad 89/29’.

Zoosphaerium bambusoides Wesener & Bespalova,

2010

Zoosphaerium bambusoides Wesener et al. 2010a: 140

(first description).

Types: M holotype.

Localities: Madagascar, Province Toamasina, Betampona.

Depositories: CAS BLF 13126-B, CASENT 9032797.

Zoosphaerium bilobum Wesener, 2009

Zoosphaerium bilobum Wesener, 2009: 90 (first descrip.)

Zoosphaerium bilobum :— Wesener et al. 2010a: 145 (com¬

parison).

Types: M holotype.

Localities: Madagascar, Province Toliara, Foret de Vohiba-

sia.

Depositories: FMNH-INS 5457.

Zoosphaerium coquerelianum (de Saussure & Zehntner,

1897)

Sphaerotherium coquerelianum de Saussure & Zehntner,

1897: pi. 1, figs 1-2 (first description)

Sphaerotherium coquerelianum :— de Saussure & Zehnt¬

ner 1902: 42 (description)

Zoosphaerium coquerelianum :— Jeekel 1999: 11 (new

combination)

Zoosphaerium coquerelianum :— Enghoff2003: 620 (list)

Zoosphaerium coquerelianum :— Wesener & Sierwald

2005a: 29 (comparison)

Zoosphaerium coquerelianum :— Wesener & Wagele

2008: 38 (redescription)

Zoosphaerium coquerelianum:— Wesener 2009: 131 (list)

Zoosphaerium coquerelianum:— Wesener & Vanden¬

Spiegel 2009: 548 (morphological phylogenetic analy¬

sis)

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Thomas Wesener

— Heligmasoma errans Chamberlin, 1921: 48, synony-

mized under S. coquerelianum by Wesener & Wagele

2008: 38

— Zoosphaerium amittum Chamberlin, 1921: 59, syno-

nymized under S. coquerelianum by Wesener & Wage¬

le 2008: 38.

Types: M lectotype, 3 F paralectotypes (designated by We¬

sener & Wagele 2008: 38).

Localities: Madagascar.

Depositories: 1 M lectotype, 2 F paralectotypes MNHN

CB015; 1 F paralectotype MNHN CB018.

Zoosphaerium denticulatum Wesener, 2009

Zoosphaerium denticulatum Wesener, 2009: 108 (first de¬

scription).

Types: M holotype.

Localities: Madagascar, Province Fianarantsoa, Androm-

bovata.

Depositories: FMNH-INS 5446.

Zoosphaerium ignotum Wesener, 2009

Zoosphaerium ignotum Wesener, 2009: 97 (first descrip.)

Types: M holotype, 2 F paralectotypes.

Localities: Madagascar, east coast, ‘waterfalls of ‘Uamot-

na?’

Depositories: SMF

Zoosphaerium isalo Wesener, 2009

Zoosphaerium isalo Wesener, 2009: 87 (first description)

Zoosphaerium isalo -- Wesener et al. 2010a: 145 (com¬

parison).

Types: M holotype.

Localities: Madagascar, Province Fianarantsoa, PN d’Isa¬

lo.

Depositories: CAS BLF 7301.

Zoosphaerium muscorum Wesener & Bespalova, 2010

Zoosphaerium muscorum Wesener et al. 2010a: 136 (first

description).

Types: M holotype.

Localities: Madagascar, Province Antsiranana, Anjana-

haribe-Sud.

Depositories: FMNH-INS 7898.

Zoosphaerium smaragdinum Wesener, 2009

Zoosphaerium smaragdinum Wesener, 2009: 80 (first des¬

cription)

Zoosphaerium smaragdinum.— Wesener et al. 2010a: 159

(additional specimens).

Types: M holotype, 1 F paratype.

Localities: Madagascar, PN Marojejy.

Depositories: CAS BLF 9079.

Zoosphaerium tainkintana Wesener, 2009

Zoosphaerium tainkintana Wesener, 2009: 100 (first de¬

scription).

Types: M holotype; 2 M, 3 F paratypes.

Localities: Madagascar, Province Antsiranana, Foret de

Binara.

Depositories: Holotype CAS BLF 9555; Paratypes: 3 F

CAS BLF 9555; 2 M, 3 F CAS BLF 9658.

Zoosphaerium tampolo Wesener, 2009

Zoosphaerium tampolo Wesener, 2009: 94 (first descrip.)

Zoosphaerium tampolo.— Wesener et al. 2010a: 159 (ad¬

ditional specimens).

Types: M holotype; 13 M, 5 F paratypes.

Localities: Madagascar, Province Toamasina, Tampolo.

Depositories: holotype FMNH-INS 3958; Paratypes: 2 M,

3 F FMNH-INS 3958; 8 M, 2 F FMNH-INS 4030; 3 M

FMNH-INS 4035.

Zoosphaerium tigrioculatum Wesener & Bespalova, 2010

Zoosphaerium tigrioculatum Wesener et al. 2010a: 144

(first description).

Types: M holotype.

Localities: Madagascar, Province Fianarantsoa, PN d’And-

ringitra.

Depositories: CAS BLF 13757 (CASENT 9032798).

Zoosphaerium trichordum Wesener, 2009

Zoosphaerium trichordum Wesener, 2009: 76 (first des¬

cription).

Types: M holotype, 2 F paratypes.

Localities: Madagascar, Province Antsiranana, Foret

d’Ampondrabe.

Depositories: CAS BLF 9973.

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©ZFMK

Species Catalogue Sphaerotheriida

Zoosphaerium villosum Wesener & Sierwald, 2005

Zoosphaerium villosum Wesener & Sierwald, 2005a: 19

(first description)

Zoosphaerium villosum.— Wesener & Wagele 2008: 42

(new locality)

Zoosphaerium villosum — Wesener 2009: 131 (list)

Zoosphaerium villosum — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium villosum :-- Wesener et al. 2010a: 160 (new

locality).

Types: M holotype, 2 M, 1 F paratypes.

Localities: Madagascar, SE coast, SF Tampolo (Type);

‘Tsianovoha’; Reserve Betampona; Foret Betaolana.

Depositories: Holotype FMNH-INS 3958. Paratypes: 1 M

FMNH-INS 8247; 1 F FMNH-INS 8248; 1 M CAS.

Zoosphaerium viridissimum Wesener, 2009

Zoosphaerium viridissimum Wesener, 2009: 104 (first des¬

cription)

Zoosphaerium viridissimum :— Wesener et al. 2010a: 160

(additional specimens).

Types: M holotype, 2 M, 1 F paratypes.

Localities: Madagascar, PN Marojejy.

Depositories: Holotype CAS BLF 9244. Paratypes: 1 F

CAS BLF 9244; 1 M CAS BLF 9079; 1 M CAS BLF

9325.

Zoosphaerium voeltzkowianum (de Saussure & Zehntner,

1897)

Sphaerotherium voeltzkowianum de Saussure & Zehntner,

1897: pi. 5, fig. 21 (first description)

Sphaerotherium voeltzkowianum — 1901: 435 (descrip.)

Sphaerotherium voeltzkowianum :— 1902: 52 (descript.)

Zoosphaerium voeltzkowianum :— Jeekel 1999: 12 (new

combination)

Zoosphaerium voeltzkowianum :— Enghoff2003: 620 (list)

Zoosphaerium voeltzkowianum :— Wesener & Sierwald

2005a: 29 (comparison)

Zoosphaerium voeltzkowianum :— Wesener & Wagele

2008: 42 (redescription)

Zoosphaerium voeltzkowianum :— Wesener & VandenSpie¬

gel 2009: 548 (morphological phylogenetic analysis)

— Sphaerotherium imbecillum de Saussure & Zehntner,

1897: pi. 5, fig. 20 (first description), synonymized un¬

der S. voeltzkowianum by Wesener & Wagele 2008: 42.

Zoosphaerium imbecillum :— Jeekel 1999: 12 (new com¬

bination)

Zoosphaerium imbecillum :— Enghoff 2003: 620 (list)

Zoosphaerium imbecillum — Wesener & Sierwald 2005a:

29 (comparison).

— Sphaerotherium globulus de Saussure & Zehntner,

1902: 34 (first description), synonymized under S.

voeltzkowianum by Wesener & Wagele 2008: 42. with

designation of lectotype.

Zoosphaerium globulus.—] eekel 1999: 12 (new combina¬

tion)

Zoosphaerium globulus :— Enghoff 2003: 620 (list)

Zoosphaerium globulus :— Wesener & Sierwald 2005a: 29

(comparison).

Remark: At the time of redescription (Wesener & Wagele

2008), only single male specimens of Z imbecillum and

Z globulus were found in the collections at the MNHN,

then referred to as holotypes. Because additional type

specimens were discovered at the MHNG, the male

MNHN type specimens used for the redescription are here¬

with selected as lectotypes.

Types: M lectotype, 2 F paralectotypes (designated by We¬

sener & Wagele 2008: 42).

Localities: Madagascar, Nosy Be.

Depositories: MNHN CB046.

The Zoosphaerium blandum species-group

Wesener 2009: 110

Zoosphaerium blandum (de Saussure & Zehntner, 1902)

(Fig. 14H)

Sphaerotherium blandum de Saussure & Zehntner, 1902:

Sphaerotherium (Globotherium) blandum — Demange

1969: 485 (additional locality)

Zoosphaerium blandum :— Jeekel 1999: 12 (new comb.)

Zoosphaerium blandum :— Enghoff 2003: 6202 (list)

Zoosphaerium blandum :— Wesener & Wagele 2008: 63

(redescription)

Zoosphaerium blandum — Wesener 2009: 132 (list)

Zoosphaerium blandum — Wesener & VandenSpiegel

2009: 548 (morphological phylogenetic analysis)

Zoosphaerium blandum:- Wesener et al. 2010b: 1185

(molecular phylogenetic analysis)

— Sphaerotherium stridulator de Saussure & Zehntner,

1902: 66 (first description), synonymized under S.

blandum by Wesener & Wagele 2008: 63, with lecto¬

type designation.

Zoosphaerium stridulator.— Jeekel 1999: 12 (new com¬

bination)

Zoosphaerium stridulator — Enghoff 2003: 621 (list).

Remarks: At the time of redescription (Wesener & Wagele

2008), only a single male specimen of Z blandum was

found in the collections at the MNHN, then referred to as

holotype. Because additional type specimens were discov¬

ered at the MHNG, the MNHN type specimen used for

the redescription is herewith selected as a lectotype.

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Thomas Wesener

Types: M lectotype (designated herewith), several

paratypes.

Localities: Madagascar, Province Toliara, col de

Sakavalana (Type); SE Madagascar spiny forest zone.

Depositories: Lectotype MNHN CB011, parataypes MH-

NG.

Zoosphaerium pseudoblandum Wesener, 2009

(Fig. 14G)

Zoosphaerium pseudoblandum Wesener, 2009: 110 (first

description)

Zoosphaerium pseudoblandum.— Wesener & Vanden

Spiegel 2009: 548 (morphological phylogenetic analy¬

sis)

Zoosphaerium pseudoblandum :-- Wesener et al. 2010b:

1185 (molecular phylogenetic analysis).

Types: M holotype, 3 M, 3 F paratypes.

Localities: Madagascar, Province Toliara, Andohahela,

Ebosika.

Depositories: FMNH-INS ‘W107’.

The Zoosphaerium piligerum species-group

Wesener & Wagele 2008: 57

Zoosphaerium micropiligerum Wesener, 2009

Zoosphaerium micropiligerum Wesener, 2009: 115 (first

description).

Types: 1 M holotype, 9 M, 2 F paratypes.

Localities: Madagascar, Province Toliara, Andohahela par¬

cel 1, 20 km SE Andranodambo (Type); Andohahela, Col

du Sedro.

Depositories: Holotype, 1 Mparatype, FMNH-INS 5399;

FMNH-INS 5416.

Zoosphaerium piligerum (de Saussure & Zehntner,

1897)

Sphaerotherium piligerum de Saussure & Zehntner,

1897: pi. 5, fig. 19 (first description)

Sphaerotherium piligerum :— de Saussure & Zehntner

1902: 36 (description)

Zoosphaerium piligerum.— Jeekel 1999: 11 (new comb.)

Zoosphaerium piligerum — Enghoff 2003: 620 (list)

Zoosphaerium piligerum :— Wesener & Wagele 2008: 57

(redescription)

Zoosphaerium piligerum :— Wesener 2009: 132 (list).

Remarks: At the time of redescription (Wesener & Wagele

2008), only a single male specimen of Z. piligerum was

found in the collections at the MNHN, then referred to as

holotype. Because additional type specimens were discov¬

ered at the MHNG, the MNHN type specimen used for

the redescription is herewith selected as a lectotype.

Types: M lectotype (MNHN), several paratypes.

Localities: Madagascar (Type); Province Antananarivo,

Andranomay.

Depositories: Lectotype MNHN CB040, paratypes MH¬

NG.

Zoosphaerium pseudopiligerum Wesener, 2009

Zoosphaerium pseudopiligerum Wesener, 2009: 118 (first

description)

Zoosphaerium pseudopiligerum :— Wesener et al. 2010a:

159 (additional specimens).

Types: M holotype, 5 M, 5 F paratypes.

Localities: Madagascar, Province Toliara, Andohahela par¬

cel 1, 15 km NW Eminiminy (Type); Andohahela parcel

1, 20 km SE Andranondambo.

Depositories: MNHN CB040.

Species unassigned to a species-group

Zoosphaerium subreflexum Jeekel, 1999

Zoosphaerium subreflexum Jeekel, 1999: 5 (first descrip.)

Zoosphaerium subreflexum :— Enghoff 2003: 620 (list)

Zoosphaerium subreflexum :— Wesener & Wagele 2008:

63 (list)

Zoosphaerium subreflexum :— Wesener 2009: 132 (list).

Types: M holotype, 2 F paratypes.

Localities: Madagascar, Province Fianarantsoa, Andringi-

tra.

Depositories: ZMA.

Zoosphaerium analavelona Wesener, 2009

Zoosphaerium analavelona Wesener, 2009: 122 (first de¬

scription)

Zoosphaerium analavelona — Wesener et al. 2010a: 159

(additional specimens).

Remarks: Z analavelona shares numerous telopod char¬

acters with Z fisheri Wesener, 2009 and can probably be

placed in a species-group with the latter.

Types: M holotype, 29 M, 12 F paratypes.

Localities: Madagascar, Province Toliara, Foret

Analavelona.

Depositories: Holotype, FMNH-INS 5431; Paratypes:

FMNH-INS 5431, 23 M, 11 F; FMNH-INS 5449, 3 M;

FMNH-INS 5427,1 M; FMNH-INS 5439,1 M; CAS BLF

7819, 1 M, 1 F.

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Species Catalogue Sphaerotheriida

Zoosphaerium fisheri Wesener, 2009

Zoosphaerium fisheri Wesener, 2009: 126 (first descrip¬

tion).

Types: M holotype, 3 M, 4 F paratypes.

Localities: Madagascar, Mandraka.

Depositories: SMF.

Zoosphaerium darthvaderi Wesener & Bespalova, 2010

Zoosphaerium darthvaderi Wesener et al., 2010a: 148

(first description).

Types: M holotype, 3 M paratypes.

Localities: Madagascar, Province Antananarivo, RS

d’Ambohitantely.

Depositories: FMNH-INS 7884, holotype; Paratypes:

FMNH-INS 7884; 8185; 8187.

Zoosphaerium heleios Wesener & Bespalova, 2010

Zoosphaerium heleios Wesener et al. 2010a: 152 (first des¬

cription)

Types: M holotype, 1 M, 2 F, 2 immature paratypes.

Localities: Madagascar, Province Toamasina, Torotorofot-

sy.

Depositories: CAS BLF 10673 (CASENT9032796),

holotype; CAS BLF 10673, paratypes.

Species only known from

females or immature males

Zoosphaerium anale (de Saussure & Zehntner, 1897)

Sphaerotherium anale de Saussure & Zehntner, 1897: pi.

4, fig. 4 (first description)

Sphaerotherium anale.— de Saussure & Zehntner 1902:

30 (description)

Zoosphaerium anale — Jeekel 1999: 9 (new combination)

Zoosphaerium anale — Enghoff 2003: 620 (list)

Zoosphaerium anale — Wesener & Wagele 2008: 76 (re¬

description)

Zoosphaerium anale - Wesener 2009: 132 (list).

Types: F lectotype (designated by Wesener & Wagele

2008: 76).

Localities: Madagascar, SW coast (Type).

Depositories: MNHN CB007.

Zoosphaerium fraternarium Jeekel, 1999

Zoosphaerium fraternarium Jeekel, 1999: 12 (replacement

name)

Zoosphaerium fraternarium :— Enghoff 2003: 620 (list)

Zoosphaerium fraternarium — Wesener & Wagele 2008:

71 (redescription)

Zoosphaerium fraternarium :— Wesener 2009: 132 (list)

— Sphaerotherium fraternum de Saussure & Zehntner,

1897: pi.5, fig. 10 (first description, e. p.), nomen no¬

vum by Jeekel 1999: 12. NOT Sphaerotherium frater¬

num Butler, 1872 (see Cyliosoma fraternum).

Types: F holotype.

Localities: Madagascar (Type); Province Fianarantsoa,

Foret FKongo (Fort Carnot).

Depositories: MNHN CB024.

Zoosphaerium immane (Karsch, 1881)

Sphaerotherium immane Karsch, 1881: 30 (first descrip¬

tion)

Sphaerotherium immane — Preudhomme de Borre 1884:

26 (list)

Sphaerotherium immane — de Saussure & Zehntner

1897: pi. 5, fig. 13 (drawing)

Sphaerotherium immane — de Saussure & Zehntner

1902: 48 (list)

Zoosphaerium immane — Jeekel 1999: 11 (new combina¬

tion)

Zoosphaerium immane — Enghoff 2003: 620 (list)

Zoosphaerium immane - Wesener & Wagele 2008: 73 (re¬

description)

Zoosphaerium immane — Wesener 2009: 132 (list).

Types: 2 F syntypes.

Localities: Madagascar.

Depositories: ZMB 715.

Zoosphaerium sakanum (Attems, 1910)

Sphaerotherium sakanum Attems, 1910: 85 (first descrip¬

tion)

Zoosphaerium sakanum :— Jeekel 1999: 12 (new combi¬

nation)

Zoosphaerium sakanum :— Enghoff 2003: 620 (list)

Zoosphaerium sakanum :— Wesener & Wagele 2008: 68

(redescription)

Zoosphaerium sakanum :— Wesener 2009: 132 (list).

Types: 2 M, 2 F syntypes.

Localities: Madagascar, Province Toamasina, Sakana.

Depositories: ZMB 5038; NHMW 2216.

Nomina dubia of Zoosphaerium

Zoosphaerium actaeon (White, 1859)

Zephronia actaeon White, 1859: 405 (first description)

Sphaerotherium actaeon.— Butler 1873: 176 (comments)

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Thomas Wesener

Zephronia actaeon — Preudhomme de Borre 1884: 23

(list)

Sphaerotherium actaeon.— de Saussure & Zehntner

1897: pi. 1, figs 1-lb (redescription without viewing of

holotype)

Sphaerotherium actaeon :— de Saussure & Zehntner

1902: 40 (redescription without viewing of holotype)

Sphaerotherium actaeon — Brolemann 1922 (list)

Sphaerotherium action [sic!]:— Overdorff 1993: 740

(lemur predation)

Zoosphaerium actaeon — Jeekel 1999: 10 (new combina¬

tion)

Zoosphaerium actaeon :— Enghoff 2003: 620 (list)

Zoosphaerium actaeon :— Wesener & Wagele 2008: 80

(comments).

Remarks: Zoosphaerium actaeon was considered a syn¬

onym of Sphaerotherium hippocastanum Gervais, 1847 by

Butler (1873: 176).

Types: Holotype of unknown sex.

Localities: Madagascar.

Depositories: NHML (presumably lost).

Zoosphaerium crassum (Butler, 1878)

Sphaerotherium crassum Butler, 1878: 301 (first descrip¬

tion)

Sphaerotherium crassum — Preudhomme de Borre 1884:

26 (list)

Sphaerotherium crassum :— de Saussure & Zehntner 1902:

68 (list)

Zoosphaerium crassum :— Jeekel 1999: 14 (new combi¬

nation)

Zoosphaerium crassum :— Enghoff 2003: 620 (list)

Zoosphaerium crassum :— Wesener & Wagele 2008: 81

(comments).

Types: Holotype of unknown sex.

Localities: Madagascar, Fianarantsoa.

Depositories: NHML (presumably lost).

Zoosphaerium elegans (Lenz, 1880)

Sphaerotherium elegans Lenz, 1880: 153 (first descrip¬

tion)

Sphaerotherium elegans :— Preudhomme de Borre 1884:

26 (list)

Sphaerotherium elegans :— de Saussure & Zehntner 1902:

48 (list)

Zoosphaerium elegans — Jeekel 1999: 14 (new combina¬

tion)

Zoosphaerium elegans :— Enghoff 2003: 620 (list)

Zoosphaerium elegans :— Wesener & Wagele 2008: 81

(comments).

Remarks: Lenz stated in a footnote that Z. elegans might

represent the undescribed male of Z hippocastanum (Ger¬

vais, 1847), and that it should be synonymized under that

name.

Types: M holotype.

Localities: Madagascar, Nosy Be.

Depositories: SMF (presumably lost).

Zoosphaerium glabrum (Butler, 1873)

Sphaerotherium glabrum Butler, 1873: 173 (first descrip¬

tion)

Sphaerotherium glabrum — Preudhomme de Borre 1884:

26 (list)

Sphaerotherium glabrum — de Saussure & Zehntner 1897:

pi. 4, fig. 3 (reprint)

Sphaerotherium glabrum :— de Saussure & Zehntner 1902:

34 (list)

Zoosphaerium glabrum — Jeekel 1999: 9 (new comb.)

Zoosphaerium glabrum :— Enghoff 2003: 620 (list)

Zoosphaerium glabrum :— Wesener & Wagele 2008: 82

(comments).

Types: Two syntypes of unknown sex.

Localities: Madagascar.

Depositories: NHML (presumably lost).

Zoosphaerium hippocastanum (Gervais, 1847)

Sphaerotherium hippocastanum Gervais, 1847: 83 (first

description)

Sphaerotherium hippocastanum :— Butler 1873: 176

(comment)

Sphaerotherium hippocastanum :— Lenz 1880: 153 (com¬

ment)

Sphaerotherium hippocastanum :— Preudhomme de Borre

1884: 26 (list)

Sphaerotherium hippocastanum :— Brancsik 1892: 258

(new record)

Zoosphaerium hippocastanum :— Jeekel 1999: 10 (new

combination)

Zoosphaerium hippocastanum :— Enghoff2003: 620 (list)

Zoosphaerium hippocastanum :— Wesener & Wagele

2008: 80 (redescription).

Remarks: The 160-year old holotype of Z hippocastanum

is completely fragmented. The specimens determined by

de Saussure and Zehntner as Z hippocastanum and used

for illustrations do include more than one species, the spec¬

imen used for illustrations represents in fact Zoosphaeri¬

um neptunus (Butler, 1872). The material used for re¬

description by Brolemann (1922) also differs from the

holotype of Z hippocastanum , and currently represents the

type series of Z broelemanni Wesener, 2009.

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Species Catalogue Sphaerotheriida

Types: F holotype (completely fragmented).

Localities: Madagascar, Nosy Be.

Depositories: MNHN CB027.

Zoosphaerium lamprinum (Butler, 1878)

Sphaerotherium lamprinum Butler, 1878: 300 (first de¬

scription)

Sphaerotherium lamprimum [sic!]:— Preudhomme de

Borre 1884: 27 (list)

Sphaerotherium lamprinum.— de Saussure & Zehntner

1897: pi. 5, figs 17-17b (drawing, mis identification?)

Sphaerotherium lamprinum :— de Saussure & Zehntner

1902: 38 (list, mis identification?)

Zoosphaerium lamprinum :— Jeekel 1999: 11 (new com¬

bination)

Zoosphaerium lamprinum :— Enghoff 2003: 620 (list)

Zoosphaerium lamprinum :— Wesener & Wagele 2008: 82

(comments).

Remarks: All specimens used for redescription by de Saus¬

sure & Zehntner (1902) represent young immatures, un¬

like the original Z. lamprinum. The drawing provided by

de Saussure and Zehntner clearly shows a female of

Zoosphaerium neptunus.

Types: Holotype of unknown sex.

Localities: Madagascar, Fianarantsoa.

Depositories: NHML (presumably lost).

Zoosphaerium latum (Butler, 1872)

Sphaerotherium latum Butler, 1872: 358 (first description)

Sphaerotherium latum — Butler 1873: 176 (list)

Sphaerotherium latum — Preudhomme de Borre 1884: 27

(list)

Sphaerotherium latum — de Saussure & Zehntner 1897:

pi. 5, fig. 14 (drawing)

Sphaerotherium latum — de Saussure & Zehntner 1902:

49 (list)

Zoosphaerium latum — Jeekel 1999: 11 (new combination)

Zoosphaerium latum — Enghoff 2003: 620 (list)

Zoosphaerium latum — Wesener & Wagele 2008: 82

(comments).

Remarks: The specimens used by de Saussure and Zehnt¬

ner (1897/1902) most likely do not represent the true Z.

latum , but Z neptunus.

Types: Holotype of unknown sex.

Localities: Madagascar, northern part.

Depositories: NHML (presumably lost).

Zoosphaerium reflexum (Brolemann, 1922)

Sphaerotherium ( Globotherium ) reflexum Brolemann,

1922: 238 (first description)

Zoosphaerium reflexum :— Jeekel 1999: 13 (new combi¬

nation)

Zoosphaerium reflexum :— Enghoff 2003: 620 (list)

Zoosphaerium reflexum — Wesener & Wagele 2008: 81

(comments).

Types: F holotype.

Localities: Madagascar, Antananarivo.

Depositories: Academie malgache (presumably lost).

Zoosphaerium ? reticulatum (Butler, 1878)

Sphaerotherium reticulatum Butler, 1878: 301 (first de¬

scription)

Sphaerotherium reticulatum :— Preudhomme de Borre

1884: 27 (list)

Sphaerotherium reticulatum :— de Saussure & Zehntner

1902: 68 (list)

Zoosphaerium reticulatum :— Jeekel 1999: 14 (new com¬

bination)

Zoosphaerium reticulatum — Enghoff 2003: 620 (list)

Zoosphaerium reticulatum :— Wesener & Wagele 2008: 78

(redescription).

Remarks: All characters, except the antennae, that would

allow for a placement of the species into a genus to be

made are missing from the types.

Types: 2 syntypes of unknown sex.

Localities: Madagascar.

Depositories: NHML BM1877-10.

Zoosphaerium stigmaticum (Butler, 1873)

Sphaerotherium stigmaticum Butler, 1873: 178, pi. 19, fig.

3 (first description)

Sphaerotherium stigmaticum :— Preudhomme de Borre

1884: 28 (list)

Sphaerotherium stigmaticum :— de Saussure & Zehntner

1897: pi. 5, fig. 18 (reprint of Butler’s figure)

Sphaerotherium stigmaticum :— de Saussure & Zehntner

1902: 67 (list)

Zoosphaerium stigmaticum:— Jeekel 1999: 14 new comb.)

Zoosphaerium stigmaticum — Enghoff 2003: 620 (list)

Zoosphaerium stigmaticum :— Wesener & Wagele 2008:

81 (comments).

Remarks: No male Sphaerotheriida from Madagascar has

a knob on the anal shield as displayed in the drawings by

Butler. However, such a knob is common in the Australian

genus Cyliosoma (Wesener 2014).

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Thomas Wesener

Types: 2 syntypes of unknown sex.

Localities: Madagascar.

Depositories: NHML (presumably lost).

Zoosphaeriuml testaceum (Olivier, 1792)

Julus testaceus Olivier, 1792: 414 (first description)

Zephronia testaceus.— Gervais 1837: 43 (list)

Sphaerotherium testaceus :-- Brandt 1841: 364 (list)

Zephronia testaceus — Gervais 1847: 86 (list)

Zephronia testaceus :-- Butler 1873: 179 (list)

Zephronia testacea — Preudhomme de Borre 1884: 25

(list)

Sphaerotherium testaceus — de Saussure & Zehntner

1902: 69 (list)

Zoosphaerium testaceum — Jeekel 1999: 15 (new comb.

Zoosphaerium testaceum — Enghoff 2003: 620 (list)

Zoosphaerium testaceum — Wesener & Wagele 2008: 82

(comments).

Remarks: The first description is not suitable for species

identification and the type specimens are presumably lost.

The generic determination is only based on the locality

given in the first

description.

Types: unknown.

Localities: Madagascar.

Depositories: unknown.

Family Cvliosomatidae Jeekel. 1974

Wesener 2014a: 199 (raised to family rank)

Sub tribe Cyliosomatini Jeekel, 1974: 45; Jamieson 1987:

64 (sperm morphology).

Genera included

Cyliosoma Pocock, 1895

Cynotelopus Jeekel, 1986

Type genus: Cyliosoma

20 species

Cyliosoma Pocock. 1895

Cyliosoma Pocock, 1895: 414 (first description); Brole-

mann 1914: 77 (review of genus); Silvestri 1917: 63 (re¬

description), 68 (defined as subgenus of Cyliosoma in

key); Verhoeff 1924: 42 (discussion); Attems 1926: 120

(list); Attems 1928: 203 (list); Verhoeff 1928: 80 (list); Ver¬

hoeff 1932: 1379 (list); Jeekel 1971: 24 (list); Jeekel 1974:

45 (classification); Hoffman 1980: 63 (list); Jeekel 1986:

36 (synonymized with Epicyliosoma)] Jamieson 1987: 64

(sperm morphology); Shelley et al. 2000: 12 (list); We¬

sener 2014a: 199 (redescription)

Synonyms:

— Epicyliosoma Silvestri, 1917: 68 (first description as

subgenus of Cyliosoma ), synonymized with Cyliosoma

by Wesener 2014. Jeekel 1971: 24 (list); Jeekel 1974:

45 (classification); Hoffman 1980: 63 (list); Jeekel 1986:

36 (with Cyliosoma as synonym); Shelley et al. 2000:

12 (list); Richter et al. 2002: 351 (mandible); Moir &

Harvey 2008: 113 (new species); Wesener & Vanden-

Spiegel 2009: 566 (classification); Wesener et al. 2010b:

1185 (molecular phylogenetic analysis).

Type species : Zephronia albertisii Silvestri, 1895, by sub¬

sequent designation by Jeekel 1971: 24.

— Paracyliosoma Verhoeff, 1928: 82 (first description

as subgenus of Epicyliosoma] see above), synonymized

under Epicyliosoma by Jeekel 1986: 36. Verhoeff 1932:

1383 (list); Jeekel 1971: 27 (list); Jeekel 1974: 45 (raised

to genus); Jeekel 1986: 36 (synonymized under

Epicyliosoma , described as subgenus of Epicyliosoma );

Hoffman 1980: 63 (list); Shelley etal. 2000: 12 (list).

Type species . Cyliosoma penicilligerum Verhoeff) 1928,

by monotypy.

Description: Male anal shield without large groove, on¬

ly a special field of setae present. Telopoditomere 3 of pos¬

terior telopods long and curved.

Type species: Sphaerotherium angulatum Butler, 1878

(Australia).

Distribution: Australia

Included species (19)

Cyliosoma angulatum (Butler, 1878) [Qld]

Cyliosoma denticulatum Verhoeff 1924 [Qld]

Cyliosoma excavatum Verhoeff, 1928 [NSW]

Cyliosoma fraternum (Butler, 1872) [Vic]

Cyliosoma froggatti Silvestri, 1917 [NSW]

Cyliosoma kurandanum Chamberlin, 1920 [Qld]

Cyliosoma larvalis (Butler, 1878) [torres Strait, Qld]

Cyliosoma marginepunctatum (Karsch, 1881) [Qld]

Cyliosoma mjoebergi Verhoeff, 1924 [Qld]

Cyliosoma pachygon Chamberlin, 1920 [Qld]

Cyliosoma penicilligerum Verhoeff, 1928 [NSW]

Cyliosoma penrithense Brolemann, 1913 [NSW]

Cyliosoma queenslandiae Brolemann, 1913 [Qld]

Cyliosoma queenslandicum Verhoeff, 1924 [NSW, Qld]

Cyliosoma sarahae (Moir & Harvey, 2008) [WA]

Cyliosoma sennae Silvestri, 1898 [Qld]

Cyliosoma sjoestedti Silvestri, 1917 [Qld]

Cyliosoma targionii Silvestri, 1898 [Qld]

Cyliosoma unicolor Silvestri, 1897 [Qld]

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Species Catalogue Sphaerotheriida

Cyliosoma angulatum (Butler, 1878)

Sphaerotherium angulatum Butler, 1878: 299 (first de¬

scription)

Sphaerotherium angulatum.— Preudhomme de Borre

1884: 26 (list)

Sphaerotherium angulatum :-- Pocock, 1893: 130 (new lo¬

cality)

Cyliosoma angulatum :— Pocock 1895: 414 (new combi¬

nation; designated as type species of Cyliosoma)

Sphaerotherium angulatum :— Brolemann 1913: 80 (list)

Cyliosoma angulatum.— Attems 1914: 141 (list)

Sphaerotherium angulatum :— Silvestri 1917: 61 (list)

Cyliosoma angulatum :— Chamberlin 1920: 88 (list)

Sphaerotherium angulatum :— Jeekel 1971: 24 (list)

Cyliosoma angulatum :— Jeekel 1981: 24 (list)

Sphaerotherium angulatum — Jeekel 1986: 36 (list)

Cyliosoma angulatum — Wesener 2014a: 205 (redescrip¬

tion, morphological phylogenetic analysis).

Types: F lectotype (designated by Wesener 2014)

Locality: Rockhampton, Qld.

Deposited: NHML BM77-36.

Cyliosoma denticulatum Verhoeff, 1924

Cyliosoma denticulatum Verhoeff, 1924: 45 (first descrip.)

Cyliosoma denticulatum.— Verhoeff 1928: 81 (list)

Cyliosoma denticulatum :— Jeekel 1981: 26 (list)

Epicyliosoma (Paracyliosoma) denticulatum Jeekel 1986:

37 (new combination)

Cyliosoma denticulatum — Wesener 2014a: 199 (list).

Types: 2 M, 1 F syntypes.

Locality: Blackall Range, Qld.

Deposited: NHRM?

Cyliosoma excavatum Verhoeff, 1928

Cyliosoma excavatum Verhoeff, 1928: 80 (first descrip.)

Cyliosoma excavatum :— Moritz & Fischer 1974: 369 (list)

Cyliosoma excavatum :— Jeekel 1981: 26 (list)

Epicyliosoma (Epicyliosoma) excavatum — Jeekel 1986:

37 (new combination)

Cyliosoma excavatum :— Wesener 2014a: 199 (list).

Remark: Jeekel viewed C. excavatum as quite similar to

C. froggatti, both coming from the same type locality

(Jeekel 1986: 37).

Types: Syntypes.

Locality: Upper Richmond River [NSW], unknown local¬

ity.

Deposited: AM; KS 76511; 1 F, ZMB 5744 (Moritz & Fis¬

cher 1974: 369).

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Cyliosoma fraternum (Butler, 1872)

Sphaerotherium fraternum Butler, 1872: 359 (first descrip¬

tion)

Sphaerotherium fraternum :— Butler 1873: 177 (in species

list)

Sphaerotherium fraternum — Preudhomme de Borre

1884: 26 (list)

Sphaerotherium fraternum — Attems 1914: 140 (list)

Sphaerotherium fraternum :— Silvestri 1917: 61 (list, ‘=?

Cyliosoma fraternum ’)

Cyliosoma (?) fraternum — Chamberlin 1920: 88 (new

combination)

‘Sphaerotherium’fraternum :— Jeekel 1981: 27 (list)

Sphaerotherium fraternum — Jeekel 1986: 39 (list)

Sphaerotherium fraternum :— Jeekel 1999: 12. (list)

NOT Sphaerotherium fraternum :— de Saussure & Zehnt-

ner, 1897 (= Zoosphaerium fraternarium Jeekel 1999:

12 )

Cyliosoma fraternum- Wesener 2014a: 208 (new comb.).

Type: F holotype

Locality: Victoria.

Deposited: NHML BM1860.38 (broken).

Cyliosoma froggatti Silvestri, 1917

Cyliosoma (Cyliosoma) Froggatti Silvestri, 1917: 68 (first

description)

Cyliosoma froggatti — Chamberlin 1920: 87 (list)

Cyliosoma froggatti :— Jeekel 1981: 26 (list)

Epicyliosoma (Epicyliosoma) froggatti :— Jeekel 1986: 37

(new combination)

Cyliosoma froggatti :— Wesener 2014a: 199 (list).

Remark: See remarks at C. excavatum.

Types: M and F syntypes.

Locality: Richmond River, NSW.

Deposited: Portici?

Cyliosoma kurandanum Chamberlin, 1920

Cyliosoma kurandanum Chamberlin, 1920: 85 (First de¬

scription)

Cyliosoma kurandanum — Jeekel 1981: 24 (list)

Epicyliosoma kurandanum :— Jeekel 1986: 37 (new com¬

bination)

Cyliosoma kurandanum — Wesener 2014a: 199 (list).

Type: F holotype

Locality: Kuranda, Qld.

Deposited: MCZ 4696.

©ZFMK

Thomas Wesener

Cyliosoma larvalis (Butler, 1878), new combination

Zephronia larvalis Butler, 1878: 301 (first description)

Zephronia larvalis -- Preudhomme de Borre 1884: 24 (list)

Zephronia larvalis — Attems 1914: 146 (list)

Zephronia (?) larvalis -- Chamberlin 1920: 91 (list)

‘Zephronia larvalis’.— Jeekel 1981: 24 (list)

Zephronia larvalis — Jeekel 1986: 39 (list)

Zephronia larvalis — Wesener 2014b: 364 (redescription)

— Zephronia Albertisii Silvestri, 1895a: 620 (in species

list), 635 (first description), synonymized under C. lar¬

valis by Wesener 2014b: 39

Cyliosoma (Epicyliosoma) Albertisii:— Silvestri 1917: 62

(new combination, redescription)

Cyliosoma (Epicyliosoma) albertisi:— Chamberlin 1920:

88 (list)

Zephronia albertisii:— Jeekel 1971: 24 (list, designated

type of Epicyliosoma.)

Cyliosoma albertisii Jeekel 1981: 24 (list)

Epicyliosoma (Epicyliosoma) albertisii Jeekel 1986: 36

(list, new combination)

Cyliosoma albertisii Wesener 2014a: 199 (list)

Cyliosoma albertisii Wesener 2014b: 364 (syn¬

onymized).

Types: F lectotype .

Locality: Torres Strait.

Deposited: NHML 77-29 (Butler 1878: 299).

Cyliosoma marginepunctatum (Karsch, 1881)

Sphaerotherium marginepunctatum Karsch, 1881: 28 (first

description)

Sphaerotherium marginepunctatum Preudhomme de

Borre 1884: 27 (list)

Sphaerotherium marginepunctatum:— Attems 1914: 141

(list)

Sphaerotherium marginepunctatum:— Silvestri 1917: 62

(list ‘=? Cyliosoma marginepunctatum'’)

Sphaerotherium marginepunctatum:— Moritz & Fischer

1973: 369 (list)

‘Sphaerotherium’ marginepunctatum:— Jeekel 1981: 24

(list)

Sphaerotherium marginepunctatum:— Jeekel 1986: 39

(list).

Cyliosoma marginepunctatum:— Wesener 2014a: 208

(new combination).

Type: F holotype.

Locality: Rockhampton, Qld.

Deposited: ZMB 396.

Cyliosoma mjoebergi Verhoeff, 1924

Cyliosoma queenslandicum mjobergi Verhoeff, 1924: 45

(as subspecies of‘C. queenslandicum'’)

Cyliosoma queenslandicum mjobergi:— Verhoeff 1928: 80

(list)

Cyliosoma queenslandiae mjoebergi:— Jeekel 1981: 24

(list)

Epicyliosoma (Paracyliosoma) mjoebergi:— Jeekel 1986:

37 (raised to species, new combination)

Cyliosoma mjoebergi — Wesener 2014a: 199 (list).

Remark: C. mjoebergi was described as a subspecies of

E. queenslandicum. According to Jeekel (1986: 37), C.

mjoebergi is close to, or even identical with C. queens¬

landiae.

Types: 1 M, 6 F syntypes.

Locality: Colosseum, Qld (Verhoeff 1924: 47).

Deposited: NHRM?

Cyliosoma pachygon Chamberlin, 1920

Cyliosomapachygon Chamberlin 1920: 86 (first descrip.)

Cyliosoma pachygon:— Jeekel 1981: 24 (list)

Epicyliosoma pachygon:— Jeekel 1986: 37 (new comb.)

Cyliosoma pachygon:— Wesener 2014a: 199 (list).

Type: M holotype.

Locality: Cooktown, Qld.

Deposited: MCZ 4697.

Cyliosoma penicilligerum Verhoeff, 1928

Cyliosoma (Paracyliosoma) penicilligerum Verhoeff,

1928: 80 (first description)

Cyliosoma penicilligerum:— Jeekel 1971: 27 (list)

Cyliosoma penicilligerum:— Jeekel 1981: 26 (list)

Epicyliosoma (Paracyliosoma) penicilligerum:— Jeekel

1986: 36 (new combination)

Cyliosoma penicilligerum:— Wesener 2014a: 199 (list).

Types: 4 M, 1 F syntypes.

Locality: North Dorrigo [NSW]

Deposited: AM KS76522.

Cyliosoma penrithense Brolemann, 1913

Cyliosoma penrithensis Brolemann, 1913: 85 (first de¬

scription)

Cyliosoma penrithensis:— Attems 1914: 64, 142 (list)

Cyliosoma penrithensis:— Silvestri 1917: 62 (syn¬

onymized with C. sennae)

Cyliosoma penrithensis:— Chamberlin 1920: 88 (list as

synonym of Cyliosoma sennae)

Cyliosoma penrithense — Verhoeff 1928: 8 (list)

Cyliosoma penrithense:— Jeekel 1981: 26 (list)

Epicyliosoma (Epicyliosoma)penrithense:— Jeekel 1986:

37 (new combination)

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Species Catalogue Sphaerotheriida

Cyliosoma penrithense.— Wesener 2014a: 199 (list).

Remark: Silvestri (1917) synonymized E. penrithense with

E. sennae , something not followed by Jeekel (1986: 36),

based on the widely separate type localities and differences

in the first descriptions.

Types: 1 M, 3 F syntypes.

Locality: Penrith & Camberra

Deposited. AM.

Cyliosoma queenslandiae Brolemann, 1913

Cyliosoma queenslandiae Brolemann, 1913: 80 (first de¬

scription)

Cyliosoma Queenslandiae.— Attems 1914: 142 (list)

Cyliosoma Queenslandiae :-- Silvestri 1917: 62 (syn¬

onymized with Cyliosoma unicolor )

Cyliosoma queenslandiae — Chamberlin 1920: 87 (list as

synonym)

Cyliosoma queenslandiae :— Verhoeff 1924: 45 (re¬

described as ‘ Cyliosoma queenslandicum Brol. (=

queenslandiae Brol.)’)

Cyliosoma queenslandiae :— Jeekel 1981: 24 (list)

Epicyliosoma (Paracyliosoma) queenslandiae :— Jeekel

1986: 37 (new combination)

Cyliosoma queenslandiae :— Wesener 2014a: 199 (list).

Remarks: Silvestri (1917) synonymized E. queenslandi¬

ae with E. unicolor. Both were described from the same

type locality and they are morphologically similar. Jeekel

treated both as separate species based on the very long

prefemoral process of the anterior telopods in C. unicol¬

or (Jeekel 1986: 37). Arestudy of the type material of both

species would be necessary to clarify this question.

Types: 4 M, 3 F syntypes.

Locality: Gayndah, Qld.

Deposited: AM.

Cyliosoma queenslandicum Verhoeff, 1924)

Cyliosoma queenslandicum Verhoeff, 1924: 45 (first de¬

scription as ‘ Cyliosoma queenslandicum Brol. (=

queenslandiae Brol.), from new material)

Cyliosoma queenslandicum :— Verhoeff 1928: 80 (redescr.)

Epicyliosoma (Paracyliosoma) queenslandicum :— Jeekel

1986: 37 (new combination, treated as species separate

from C. queenslandiae Brol.)

Cyliosoma queenslandicum :— Wesener 2014a: 199 (list).

Remarks: Verhoeff viewed E. queenslandicum as identi¬

cal with E. targionii. Jeekel wrote (1986: 37): “It seems

likely, however, that Verhoeff mixed up two species un¬

der the name: one from northern Queensland and one from

southern Queensland, subsequently reported also from

northern New South Wales. A restudy of the type materi¬

al, involving also the selection of a lectotype, is needed

to clarify this point.”

Types: Several M and F syntypes.

Localities: Herberton, Mt Tamborine and Malanda, Qld.

Deposited: NHRM?

Cyliosoma sarahae (Moir & Harvey, 2008)

(Figs 15A, B)

Epicyliosoma (Epicyliosoma) sarahae Moir & Harvey,

2008: 114 (first description)

Epicyliosoma sarahae - Moir et al. 2009: 1962 (Ecolo¬

gy, distribution)

Cyliosoma sarahae — Wesener 2014a: 199 (list).

Types: M holotype, 5 M, 8 F paratypes.

Locality: Cape Le Grand National Park, inland from This¬

tle Beach, WA, 34°00’20”S, 122°1L39”E, in soil.

Deposited: WAM T78757, T78948.

Cyliosoma sennae Silvestri, 1898

Cyliosoma Sennae Silvestri, 1898: 227 (first description)

Cyliosoma sennae — Brolemann 1913: 80 (list)

Cyliosoma Sennae — Attems 1914: 142 (list)

Cyliosoma (Cyliosoma) Sennae — Silvestri 1917: 62 (re¬

description)

Cyliosoma sennae :— Chamberlin 1920: 88 (list)

Cyliosoma sennae.— Jeekel 1981: 24 (list)

Epicyliosoma (Epicyliosoma) sennae :— Jeekel 1986: 37

(new combination)

Cyliosoma sennae — Wesener 2014a: 199 (list).

Remark: Silvestri (1917) viewed E. penrithense as syn¬

onymous with E. sennae. Jeekel (1986: 37) did not fol¬

low this synonymization, treating both as separate enti¬

ties based on the differences in the first descriptions and

the widely separated type localities.

Types: M and F syntypes.

Locality: Caims, Qld.

Deposited: MZUF.

Cyliosoma sjoestedti Silvestri, 1917

Cyliosoma (Epicyliosoma) Sjostedti Silvestri, 1917: 68

(first description)

Cyliosoma sjostedti — Chamberlin 1920: 88 (list)

Cyliosoma sjoestedti — Jeekel 1981: 24 (list)

Epicyliosoma sjoestedti :— Jeekel 1986: 37 (new comb.)

Cyliosoma sjoestedti :— Wesener 2014a: 199 (list).

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Thomas Wesener

Types: F holotype or syntypes.

Locality: Cardwell, Qld.

Deposited: NHRM.

Cyliosoma targionii Silvestri, 1898)

Cyliosoma Targionii Silvestri, 1898: 226 (first description)

Cyliosoma targionii.— Brolemann 1913: 80 (list)

Cyliosoma Targionii.— Attems 1914: 142 (list)

Cyliosoma (Cyliosoma) Targionii :-- Silvestri 1917: 62 (re¬

description)

Cyliosoma targionii — Chamberlin 1920: 87 (list)

Cyliosoma targionii — Jeekel 1981: 24. (list)

Epicyliosoma (Paracyliosoma) targionii — Jeekel 1986:

37 (new combination)

Cyliosoma targionii — Wesener 2014a: 199 (list).

Remarks: See E. queenslandicum for a history of the con¬

fusion of the two species.

Types: M and F syntypes.

Locality: Cairns, Qld.

Deposited: MZUF.

Cyliosoma unicolor Silvestri, 1897

Cyliosoma imicolor Silvestri, 1897a: 16 (first description)

Cyliosoma imicolor .— Brolemann 1913: 80 (list).

Cyliosoma unicolor .— Attems 1914: 142. (list)

Cyliosoma (Cyliosoma) unicolor :— Silvestri 1917: 62 (re¬

description)

Cyliosoma unicolor— Chamberlin 1920: 87 (list)

Cyliosoma unicolor— Jeekel 1981: 24. (list)

Epicyliosoma (Paracyliosoma) unicolor — Jeekel 1986: 37

(new combination)

Cyliosoma imicolor— Wesener 2014a: 199 (list).

Remarks: See remarks under C. queenslandiae above.

Types: M and F syntypes.

Locality: Gayndah, Qld.

Deposited: MTD 283.

Cynotelopus Jeekel. 1986

Cynotelopus Jeekel, 1986: 39 (first description); Shelley

et al. 2000: 12 (list, under family Sphaeropoeidae); We¬

sener & VandenSpiegel 2009: 566 (morphological phylo¬

genetic analysis); Wesener 2014a: 209 (morphological

phylogenetic analysis).

Description: Male anal shield heavily modified, with a

gigantic groove. Telopoditomere 3 of posterior telopods

wide and short.

Bonn zoological Bulletin - Supplementum 63: 1-104

Type species: Cynotelopus notabilis Jeekel, 1986, by orig¬

inal designation.

Distribution: Australia, WA.

Included species:

Cynotelopus notabilis Jeekel, 1986 [WA]

Monotypic

Cynotelopus notabilis Jeekel, 1986

(Fig. 15B)

Cynotelopus notabilis Jeekel, 1986: 40 (first description)

Cynotelopus notabilis :— Main et al. 2002: 383 (new lo¬

cality)

Cynotelopus notabilis :— Moir et al. 2009: 1962 (ecology).

Types: M holotype, 2 F paratypes.

Locality: Nomalup, WA.

Deposited: Holotype SAM OM1215; paratypes SAM.

Nomina dubia of Sphaerotheriida

Sphaerotherium convexum C.L. Koch, 1847

Sphaerotherium convexum C.L. Koch, 1847: 100 (first de¬

scription)

Sphaerotherium convexum :— C.L. Koch 1863: 31 (re¬

description)

Sphaerotherium convexum :— Butler 1873: 176 (list)

Sphaerotherium convexum :— Karsch 1881: 28 (list)

Sphaerotherium convexum — Preudhomme de Borre

1884: 26 (list)

Sphaerotherium convexum — Attems 1914: 140 (list)

Sphaerotherium convexum :— Silvestri 1917: 61 (list)

Cyliosoma (?) convexum — Chamberlin 1920: 88 (list)

Sphaerotherium convexum :— Jeekel 1981: 24 (list)

Sphaerotherium convexum :— Jeekel 1986: 39 (list).

Types: Unknown.

Locality: ‘Neu Holland.’

Deposited: Unknown.

Zephronia glaberrima Attems, 1898

Zephronia glaberrima Attems 1898: 510 (first description)

Zephronia glaberrima — Attems 1914: 145 (list)

‘Zephronia glaberrima:— Jeekel 1981: 24 (list)

Zephronia glaberrima — Jeekel 1986: 39 (list).

Types: F holotype

Locality: Cooktown, Qld.

Deposited: Unknown.

©ZFMK

Species Catalogue Sphaerotheriida

Sphaerotherium gronovii Brandt 1841

Sphaerotherium gronovii Brandt 1841a: 180

Sphaerotherium gronovii.— Butler 1873: 178 (list)

Sphaerotherium ? gronovii :— Preudhomme de Borre 1884:

26 (list).

Types: Unknown.

Locality: ‘North Sea between Norway and Great Britain’,

unknown

Deposited: Should have been in ZISP, but presumably lost.

Sphaerotherium kutorgae Brandt, 1841

Sphaerotherium kutorgae Brandt, 1841a: 176 (first de¬

scription)

Sphaerotherium kutorgae :— Butler 1873: 175 (list)

Sphaerotherium kutorgae :— Preudhomme de Borre 1884:

27 (list).

Types: Holotype of unknown sex.

Locality: Unknown

Deposited: Should have been in ZISP, but presumably lost.

Sphaerotherium tit anus Brandt, 1841

Sphaerotherium titanus Brandt, 1841a: 176 (first descrip¬

tion)

Sphaerotherium titanus :— Butler 1873: 177 (list)

Sphaerotherium titanus — Preudhomme de Borre 1884: 28

(list).

Types: Unknown.

Locality: Unknown.

Deposited: Should have been in ZISP, but presumably lost.

Nomina nuda of Sphaerotheriida

Remarks: The here listed genus names are of no status in

nomenclature, as they were not proposed in concordance

with the then valid code.

Ceylonhelum Verhoeff, 1937

Ceylonhelum — Verhoeff 1937: 81 (mention)

Ceylonhelum :— Jeekel 1971: 24 (explanation)

Remark: No type species was designated and the proposed

manuscript in which this genus was supposed to be de¬

scribed was never finished or published.

Dekanohelum Verhoeff) 1937

Dekanohelum — Verhoeff 1937: 81 (mention)

Dekanohelum — Jeekel 1971: 24 (explanation)

Remark: No type species was designated and the proposed

manuscript in which this genus was supposed to be de¬

scribed was never published.

Plethoconium Attems, 1943

Plethoconium Attems, 1943: 60 (description as subgenus)

Plethozonium [sic!]:-Attems 1953: 154 (list)

Plethoconium — Jeekel 1971: 27 (explanation)

Remark: Proposed without any species assigned to the

name and therefore not available

Rhopalopaeus Verhoeff, 1942

Rhopalopaeus :— Takakuwa 1942: 362 (mention)

Rhopalopaeus :— Jeekel 1971: 28 (explanation)

Remark: Takakuwa lists the genus name with reference

to Verhoeff. However, Verhoeff never published a paper

with such a generic name, therefore the name is not ava¬

ilable

Acknowledgements. I am indebted to many colleagues and

friends who hosted me during a visit, loaned out or collected im¬

portant specimens: First at all my fellow giant pill-millipede co¬

workers Didier van den Spiegel (MRAC) and Nattarin Wongth-

amwanich (CUMZ); the late Richard Hoffman (VMNH); Petra

Sierwald and Steve Goodman (FMNH); Jean-Jacques Geoffroy,

Mark Judson, Monique Nguyen Duy-Jacquemin, and Jean-Paul

Mauries (MNHN); Darrel Ubick, Brian Fisher and Charles Gris¬

wold (CAS); Peter Schwendinger (MHNG); Henrik Enghoff

(ZMUC); Hieronymus Dastych (ZMUH); Maurizio Mei (Roma);

Peter Jager and Julia Altmann (SMF); Jason Dunlop and Anja

Friederichs (ZMB); Janet Beccaloni (NHML); Verena Stagl, Nes-

rine Akkari and Edmund Schiller (NHMW). Many thanks to Ver¬

ena Stagl (Vienna) for sending me her draft of a type catalogue.

I thank Petra Sierwald (FMNH), Sergei Golovatch (Russian

Academy of Sciences) and Bob Mesibov (Tasmania) for read¬

ing a previous draft of the catalogue. Peter Kautt (Tubingen),

Jan Philip Oeyen (Bonn), Ame Hartig (Bielefeld), Robert Mesi¬

bov (Tasmania), Melinda Moir (Melbourne), and Michelle

Hamer (KwaZulu-Natal) allowed me to use their excellent pho¬

tographs of giant pill-millipedes. Holly E Wesener proof-read

the English.

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Thomas Wesener

INDEX TO GENERIC NAMES

(including synonyms = syn.)

Arthrosphaera 63

Borneopoeus (syn.) 42

Bothrobelum 27

Bournellum (syn.) 51

Castanotherium 42

Castanotheroides 49

Chinosphaera (syn.) 37

Cryxus 27

CyJiosoma 88

Cyliosomella (syn.) 24

Cynotelopus 92

Epicyliosoma (syn.) 88

Eubournellum { syn.) 51

Globotherium (syn.) 76

Heligmasoma (syn.) 76

Indosphaera 28

Kophosphaera 28

Kylindotherium 63

Leptotelopus 30

Leptoprotopus 38

Lissosphaera (syn.) 38

Lophozephronia (syn.) 30

Luzonosphaera (syn.) 49

INDEX TO SPECIFIC NAMES

(including synonyms = syn.)

actaeon, Zoosphaerium 85

albertisii, CyJiosoma (syn.) 90

album, Zoosphaerium 78

alluaudi, Zoosphaerium 81

alticola alticola, Zephronia 31

alticola, Sphaerotherium 52

amabile, Zoosphaerium 81

ambrense, Zoosphaerium 81

amittum, Zoosphaerium (syn.) 82

amythrum, Castanotherium 43

analavelona, Zoosphaerium 84

anale, Zoosphaerium 85

ancillare, Sphaerotherium (syn.) 54

andersoni, Procyliosoma 24

andohahela, Sphaeromimus 74

andrahomana, Sphaeromimus 74

angulatum, CyJiosoma 89

anjozorobe, Micro sphaerotherium 76

anomalum, Zoosphaerium 78

anthracina, Zephronia 31

apicale, Sphaerotherium 52

arborealis, Zoosphaerium 78

atrisparsa, Arthrosphaera 64

attemsi, Arthrosphaera 64

aureum, Zoosphaerium 78

aurivilii, Procyliosoma 25

aurocincta, Arthrosphaera 65

bambusoides, Zoosphaerium 81

banksianus, Sphaeropoeus (syn.) 39

barbata, Sphaeropoeus (syn.) 39

bengalica alticola, Zephronia 31

bicolor, Arthrosphaera 65

Microsphaerotherium

Natalobelum (syn.)

Neobournellum (syn.)

Odontosternum (syn.)

Oligaspis (syn.)

Pantitherium (syn.)

Paracyliosoma (syn.)

Prionobelum

Procyliosoma

Pulusphaera (syn.)

Rajasphaera

Sechelliosoma

Sphaerobelum

Sphaeromimus

Sphaeropoeus

Sphaerotherium

Syncyliosoma (syn.)

Tetraconosoma (syn.)

Tigridosphaera

Tonkinobelum (syn.)

Trivandrobelum (syn.)

Trochosoma (syn.)

Zephronia

Zoosphaerium

bicollis, Sphaeropoeus (syn.)

bicorne, Sphaerobelum

bilobum, Zoosphaerium

bimaculata, Tigridosphaera

blandum, Zoosphaerium

boerium, Sphaerotherium

boetonense, Castanotherium

brandti, Arthrosphaera

brevilamina, Kophosphaera

broelemanni, Zoosphaerium

butleri, Zephronia

cambodjana, Zephronia (syn.)

campanulatum, Zoosphaerium

capense, Sphaerotherium

carinata, Arthrosphaera

carinatum, Castanotherium

castaneum, Procyliosoma

castaneum, Castanotherium

celebense, Castanotherium

chitinoides, Zephronia (syn.)

cinctellum, Sphaerotherium

cinctum, Castanotherium

civicum, Sphaerotherium

clavigerum, Sphaerobelum

clivicola, Zephronia

commune, Sphaerotherium

comotti, Zephronia

compressum, Sphaerotherium

coniferum, Sphaerotherium

conspicuum, Castanotherium

convexitarsum, Sphaerotherium

convexum, Sphaerotherium

coquerelianum, Zoosphaerium

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Species Catalogue Sphaerotheriida

corrugata, Arthrosphaera 66

corystoides, Zoosphaerium 80

costatum, Castanotherium 44

craspedota, Arthrosphaera 66

crassum, Zoosphaerium 86

crepitans, Leptotelopus 30

criniceps, Castanotherium 44

crinitum, Castanotherium 44

curiosa, Indosphaera (syn.) 28

dalyi, Arthrosphaera 66

darthvaderi, Zoosphaerium 85

davisoni, Arthrosphaera 67

dawydoffi, Zephronia 31

debilis, Zephronia 32

decoratum, Castanotherium 44

delacyi delacyi, Procyliosoma 25

densipora, Zephronia 32

denticulatum, Zoosphaerium 82

denticulatum, Cyliosoma 89

dentigera, Arthrosphaera 67

devolvens, Kophosphaera 29

dicrothrix, Sphaerotherium 54

digitate, Zoosphaerium (syn.) 77

dinogonum, Sphaerotherium 54

discolour, Zoosphaerium 79

disparipora, Zephronia 32

disticta, Arthrosphaera 67

distinctum, Castanotherium 44

dollfusi, Zephronia 32

doriae, Zephronia 32

dorrigense, Procyliosma 25

dorsale, Sphaerotherium 54

dorsaloide, Sphaerotherium 55

dorsispina, Castanotherium 45

durum, Prionobelum 37

elegans, Zoosphaerium 86

elongatum, Sphaerotherium 55

endemicum, Zoosphaerium 79

eremita, Sphaerotherium 55

errans, Heligmasoma (syn.) 82

eucalyptophilum, Sphaerotherium 56

evansi, Tigridosphaera 42

everetti, Castanotherium 45

excavata, Kophosphaera 29

excavatum, Cyliosoma 89

extinctus, Sphaeropoeus 39

extraneum, Castanotherium 45

falcicorne, Castanotherium 45

feae, Indosphaera 28

feae, Zephronia 32

ferum, Castanotherium 45

fisheri, Zoosphaerium 85

floweri, Zephronia 32

forcipatum, Sechelliosoma 50

formosa, Zephronia 32

fortis, Indosphaera 28

fraternarium, Zoosphaerium 85

fraternum, Cyliosoma 89

frogatti, Cyliosoma 89

fulvicorne, Castanotherium 45

fulvum, Sphaerotherium 56

fumosa, Arthrosphaera 67

fusca, Arthrosphaera 68

gestri, Zephronia 32

giganteum, Sphaerotherium 56

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gisleni, Sphaeropoeus 39

glaberrima, Zephronia 92

glabratum, Castanotherium 45

glabrus, Sphaeropoeus 39

glabrum, Zoosphaerium 86

gladiator, Leptoprotopus 38

globulus, Zoosphaerium (syn.) 83

globusmagicus, Tigridosphaera 42

gracilis, Arthrosphaera 68

granulatum, Castanotherium 45

granulatum, Sphaerotherium 56

gronovii, Sphaerotherium 93

grossum, Sphaerotherium (syn.) 58

haackeri, Zoosphaerium 79

hainani, Prionobelum 37

hanstroemi, Sphaerotherium 56

heleios, Zoosphaerium 85

hendersoni, Arthrosphaera 68

hercules, Sphaeropoeus 39

heterostictica, Arthrosphaera 68

hippocastanum, Zoosphaerium 86

hirsti, Zephronia 33

hirsutellum, Castanotherium 46

hirsutum, Sphaerobelum 36

hirta, Zephronia 33

hosei, Castanotherium 46

humile, Castanotherium 46

hysophila, Zephronia 33

ignobile, Castanotherium 46

ignotum, Zoosphaerium 82

imbecillum, Zoosphaerium (syn.) 83

immane, Zoosphaerium 85

impuncata, Zephronia 33

indigenum, Castanotherium 46

inermis, Arthrosphaera 69

inexpectatus, Sphaeromimus 74

inferior, Zephronia 33

innominatum, Castanotherium 46

insulanum, Sphaerotherium 55

insigne, Castanotherium 46

intermedium, Sphaerotherium 57

isalo, Zoosphaerium 82

ivohibe, Sphaeromimus 74

ivohibiense, Microsphaerotherium 76

javanica, Zephronia (syn.) 46

joliveti, Prionobelum 37

juvenis, Zephronia 33

kitharistes, Sphaerotherium 57

klugii, Sphaerotherium (syn.) 53

kochii, Sphaerotherium (syn.) 59

krugeri, Sphaerotherium 57

kurandanum, Cyliosoma 89

kutorgae, Sphaerotherium 93

laeve, Castanotherium 47

laevissima, Zephronia (syn.) 34

lambertoni, Zoosphaerium 79

lamprinum, Zoosphaerium 87

lamprongse, Castanotherium 47

larvalis, Cyliosoma 90

latum, Zoosphaerium 87

lavasoa, Sphaeromimus 75

leae, Procyliosoma 25

leiosomum, Procyliosoma 26

leium, Castanotherium 47

leopardina, Arthrosphaera 69

©ZFMK

Thomas Wesener

leve, Prionobelum 37

leve, Kylindotherium 63

libidinosum, Zoosphaerium 79

lichtensteinii, Sphaerotherium 57

lignivora, Zephronia 33

lugubris, Sphaeropoeus 40

lutescens, Arthrosphaera 69

maculata, Zephronia 33

maculatus, Sphaeropoeus 40

maculosum, Prionobelum 38

madura, Arthrosphaera 70

magna, Arthrosphaera 70

magnus, Sphaeropoeus 40

mahaium, Sphaerotherium 57

majorinum, Prionobelum 38

malaccanus, Sphaeropoeus 40

mammifera, Kophosphaera 29

manca, Zephronia 33

margined a, Arthrosphaera 70

marginepunctatum, Cyliosoma 90

marmorata, Arthrosphaera 70

martensi, Kophosphaera 29

micropiligerum, Zoosphaerium 84

microstictum, Sphaerotherium (syn.) 55

millepunctatum, Sphaerotherium 58

mitoho, Zoosphaerium 79

mjoebergi, Cyliosoma 90

moderatum, Castanotherium 47

modestum, Sphaerotherium 58

modiglianii, Sphaeropoeus 40

montana, Zephronia 34

montana, Rajasphaera 50

monticola, Sphaerotherium (syn.) 53

multidentata, Prionobelum 38

muscorum, Zoosphaerium 82

musicus, Sphaeromimus 75

narcissei, Sphaerotherium 58

nebulosum, Sphaerotherium 51

nepalensis, Zephronia 34

neptunus, Zoosphaerium 11

nigriceps, Castanotherium 47

nigrinota, Zephronia 34

nigritarse, Sphaerotherium 58

nigromaculatum, Castanotherium 47

nigrum, Sphaerotherium 63

nitida, Arthrosphaera 1 1

notabilis, Cynotelopus 92

noticeps, Arthrosphaera 1 1

novae-zea/andudae, Procyliosoma 25

obscurum, Castanotherium 47

obtusum, Sphaerotherium 58

ornatum, Castanotherium 47

oval is, Cryxus 28

ovalis, Zephronia 34

pachygon, Cyliosoma 90

pellita, Zephronia 34

pellitum, Castanotherium 47

pellitus, Sphaeropoeus 40

pelloceps, Arthrosphaera 1 1

penicilligerum, Cyliosoma 90

penrithense, Cyliosoma 90

perbrincki, Sphaerotherium 58

permodestum, Sphaerotherium 59

philippinus, Castanotheroides 49

pilifera, Arthrosphaera 1 1

Bonn zoological Bulletin - Supplementum 63: 1-104

piligerum, Zoosphaerium 84

pilosum, Castanotherium 48

pinnatum, Sphaerotherium 59

plagiarium, Sphaerotherium 59

plasoni, Castanotherium 48

platylabum, Zoosphaerium 80

politissima, Kophosphaera 29

politum, Zephronia (syn.) 34

politus, Sphaeropoeus 40

porosus, Castanotheroides 50

priapus, Zoosphaerium 19

profuga, Zephronia 34

pseudoblandum, Zoosphaerium 84

pseudopiligerum, Zoosphaerium 84

pseudoplatylabum, Zoosphaerium 80

pseudopriapus, Zoosphaerium 80

pubescens, Sphaerotherium (syn.) 60

pulchellum, Zoosphaerium 80

punctatum, Sphaerotherium 59

punctipes, Oligaspis (syn.) 60

punctulatissimus, Sphaeropoeus 40

punctulatum, Sphaerotherium 59

puverea, Zephronia (syn.) 55

pygidiale, Zoosphaerium (syn.) 78

pygostolis, Arthrosphaera 71

pyrrhomelanum, Castanotherium 48

queenslandiae, Cyliosoma 91

queenslandicum, Cyliosoma 91

reflexum, Zoosphaerium 87

repulsum, Sphaerotherium (syn.) 54

reticulatum, Zoosphaerium 87

retusum, Sphaerotherium (syn.) 55

ridleyi, Zephronia 35

rotundatum, Sphaerotherium 60

rudebecki, Sphaerotherium 60

rufipes, Castanotherium 48

ruginosa, Arthrosphaera 71

rugosa, Arthrosphaera 12

rugosum, Bothrobelum 21

rugulosum, Sphaerotherium 60

saintelucei, Sphaeromimus 75

sakanum, Zoosphaerium 85

sarahae, Cyliosoma 91

scholastica, Arthrosphaera 12

selindum, Sphaerotherium 61

semilaevis, Zephronia 35

sennae, Cyliosoma 91

separatum, Sphaerobelum 36

severa, Arthrosphaera 12

shivapuri, Kophosphaera 29

siamensis, Zephronia 35

silvestrii, Arthrosphaera 12

similare, Sphaerotherium 61

simplex, Castanotherium 48

sinuatus, Castanotheroides 50

sjoestedti, Cyliosoma 91

smaragdinum, Zoosphaerium 82

solitarium, Sphaerotherium 61

solitarium, Zoosphaerium 80

sparsepunctatum, Castanotherium 48

speciosus, Sphaeropoeus 41

specularis, Zephronia 35

spinatum, Sphaerotherium 61

splendidus, Sphaeromimus 75

stellatum, Castanotherium 48

©ZFMK

Species Catalogue Sphaerotheriida

steppense, Sphaerotherium 61

stigmaticum, Zoosphaerium 87

stolli, Sphaeropoeus 41

stridula, Arthrosphaera 72

stridulator, Zoosphaerium (syn.) 83

striolatum delacyi, Procyliosoma 25

subdorsale, Sphaerotherium 61

submite, Sphaerotherium 62

subreflexum, Zoosphaerium 84

sulcatulum, Bothrobelum 27

sulcicollis, Castanotherium 48

sumatrensis, Sphaeropoeus 41

suspectum, Castanotherium 49

tainkintana, Zoosphaerium 82

tampolo, Zoosphaerium 82

targionii, Cyliosoma 92

tasmanicum, Procyliosoma 26

tatusiaeformis, Sphaeropoeus 41

tenuitarse, Sphaerotherium 62

testaceum, Zoosphaerium 88

thurstoni, Arthrosphaera 72

tigratus, Sphaeropoeus 41

tigrina, Zephronia 35

tigrinoides, Zephronia 35

tigrioculatum, Zoosphaerium 82

titanus, Sphaeromimus 75

titanus, Sphaerotherium 93

tivium, Castanotherium 49

tomentosum, Sphaerotherium 62

transitiva, Arthrosphaera 73

transzambeziacum, Sphaerotherium 62

trichopygum, Sphaerotherium (syn.) 54

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Bonn zoological Bulletin - Supplementum 63: 1-104

©ZFMK

Summary

This monograph provides the first comprehensive overview of the

current state of knowledge of the ecology, development, morphol¬

ogy, and diversity of the giant pill-millipedes, order Sphaerotheriida.

It summarizes all known aspects of the life history and ecology,

including predators and parasites, of giant pill-millipedes and maps

the biogeography of the order and its families. This work includes

illustrations, primarily based on scanning electron microscopy

images, of all major general morphological characters of the giant

pill-millipedes. To capture the intraordinal variation of giant pill-mil¬

lipedes, it further includes illustrations of the female vulva and male

posterior telopods of all known main groups and genera of the

order. In addition, the monograph provides a detailed list of all 39

known unique characters of members of the Sphaerotheriida.

The catalogue part presents the first complete list of all described

species, genera, and higher categories of the giant pill-millipedes. It

includes a key to all five Sphaerotheriida families, short descriptions

of the key characters for each of the 24 genera, and a list of all 351

valid Sphaerotheriida species with their synonyms. For each species,

the catalogue further provides the number and location of type

depositories as well as the locality data from the literature. For each

family, genus, and species it includes bibliographic citations, includ¬

ing all subsequent citations, in chronological order. The monograph

also contains a complete reference list with more than 280 entries

about the Sphaerotheriida.

Bonn zoological Bulletin - Supplementum Vol. 63 (2016)

Managing Editor: Ralph S. Peters

Zoologisches Forschungsmuseum Alexander Koenig -

Leibniz-lnstitut fur Biodiversitat der Tiere (ZFMK)

Adenaueralllee 160, D-53113 Bonn, Germany

ISBN-13: 978-3-925382-67-3

ISSN: 0302-671X

Cover illustration:

The Montagne de Frangais giant pill-millipede, Zoosphaerium solitarium Wesener, 2009, one of

Madagascar's numerous microendemic species, occuring only on top of a single karstic hill in northern

Madagascar. Photos by Jorn Kohler, Darmstadt.

Bundesministerium

furBildung

und Forschung

Ministerium fur Innovation,

Wissenschaft und Forschung

des Landes Nordrhein-Westfalen

Bonn zoological Bulletin - Supplementum Volume 63 (2016)