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Zoological Journal of the Linnean Society, 2013, 169, 737–764. With 87 figures
On the largest chelodesmid millipedes: taxonomic
review and cladistic analysis of the genus Odontopeltis
Pocock, 1894 (Diplopoda; Polydesmida; Chelodesmidae)
JOÃO P. P. PENA-BARBOSA1,2*, PETRA SIERWALD3 and ANTONIO D. BRESCOVIT2
1
Pós-graduação em Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, Brazil
Laboratório Especial de Coleções Zoológicas, Instituto Butantan, Avenida Vital Brasil, 1500,
05503-090, São Paulo, Brazil
3
Department of Zoology, Insect Division, Field Museum of Natural History, 1400 South Lake Shore
Drive, Chicago, IL 60605, USA
2
Received 16 April 2013; revised 28 August 2013; accepted for publication 28 August 2013
The chelodesmid genus Odontopeltis Pocock, 1894, is revised. It currently contains eight species, four of which are
described as new here: Odontopeltis aleijadinho sp. nov., Odontopeltis donabeja sp. nov., Odontopeltis
tiradentes sp. nov., and Odontopeltis xica sp. nov. Additional descriptive data are provided for Odontopeltis
anchisteus Hoffman, 1981, Odontopeltis clarazianus (Humbert & deSaussure, 1869), Odontopeltis giganteus
Schubart, 1849, and Odontopeltis conspersus (Perty, 1844); for the latter species a neotype is designated herein. The
current position of all species ever associated with the genus is presented. Examination of type material revealed
that the following species are not members of the genus Odontopeltis: Odontopeltis balzanii Silvestri, 1895,
Odontopeltis borellii Silvestri, 1895, Polydesmus gracilipes Humbert & deSaussure, 1870, and Odontopeltis
proxima Silvestri, 1895. The type specimen of Rhacophorus decoloratus Koch, 1847, is lost and the species
is considered incertae sedis. The terminology of gonopod structures is discussed. A phylogenetic analysis of
Odontopeltis based on 47 morphological characters, with species of the genus Odontopeltis and six outgroup taxa
is presented. Odontopeltis is monophyletic, with Rondonaria as its closest sister-group. The included members of
the tribe Telonychopodini (Pantanalodesmus, Telonychopus, and Manfrediodesmus) form a monophyletic group; the
tribe Macrocoxodesmini (Macrocoxodesmus and Eucampesmella) is paraphyletic in this data set.
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764.
doi: 10.1111/zoj.12086
ADDITIONAL KEYWORDS: Brazil – gonopod terminology – Macrocoxodesmini – Telonychopodini.
INTRODUCTION
Millipedes of the genus Odontopeltis Pocock, 1894,
comprise the largest members of the family
Chelodesmidae (Brewer, Sierwald & Bond, 2012); the
longest specimens measure up to 107 mm. Members
of the genus are conspicuous and colourful (Fig. 1).
The genus has had a tumultuous taxonomic history
and even its placement in the family Chelodesmidae
is somewhat circumstantial. As the late Hoffman
*Corresponding author. E-mail: [email protected]
(1982a) stated: ‘the family is extremely large and
diverse, and difficult to define succinctly’. The placement of Odontopeltis into the Chelodesmidae is currently not supported by well-defined apomorphies
of the family, but rather by the fact that the genus
cannot be accommodated easily in any other
polydesmid family. Hoffman (1981a) resolved the
convoluted taxonomic history of the genus name
Odontopeltis, revalidated the genus, and revised the
then-known four species of Odontopeltis. He noted
that Odontopeltis may be related to Telonychopus
Verhoeff, 1951, Leiodesmus Silvestri, 1897, and
Manfrediodesmus Schubart, 1949, and listed the
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
737
738
J. P. P. PENA-BARBOSA ET AL.
Figure 1. Live male specimen of Odontopeltis donabeja
sp. nov., from Estação Biológica de Peti, CEMIG, São
Gonçalo do Rio Abaixo, Minas Gerais, Brazil.
genus in the chelodesmine tribe Telonychopodini
(Hoffman, 1980: 151) in his monumental work Classification of the Diplopoda. In subsequent publications (Hoffman, 1981a, 2000), Odontopeltis was not
placed in any chelodesmid tribe and thus its systematic position within the Chelodesmidae remains
unresolved.
This study is a revision of all the species of the
genus, contains a redescription of the type species of
the genus, Odontopeltis conspersus (Perty, 1833), and
introduces and describes four new species in the
genus. As in other helminthomorph millipedes, the
male gonopods are complex and provide a rich suite of
characters, but employing the gonopod characters in
rigorous phylogenetic analyses is currently hampered
by a lack of primary homology hypotheses of the
various branches and apophyses displayed by these
organs, even within the family Chelodesmidae. Succinct character delimitation is also impacted by the
usage of several differing terminologies for the constituent parts; a comprehensive ontology for these
features has not been developed. This study reviews
and reconciles the gonopod terminologies employed
by previous authors for the genus, and examines the
systematic position of the genus within the large
family Chelodesmidae.
SYSTEMATIC
POSITION OF
ODONTOPELTIS
Comprising well over 5000 described species and
approximately 1400 genera arranged in 30 families,
the Polydesmida Leach, 1814, is the most species-rich
order in the Diplopoda (Brewer et al., 2012). Male
members of the order have species-specific gonopods
on the seventh body ring: the anterior leg pair of
this body ring is modified to a sperm-transferring
organ in males; the posterior legs of the same ring
are unmodified walking legs. Features in the often
complex male reproductive structures are the main
species-delimitating characters (Schubart, 1949;
Hoffman, 1980).
The family Chelodesmidae, with approximately
800 species, is currently the second most speciose
family of the order, after the Paradoxosomatidae.
The family Chelodesmidae was proposed by
Cook (1895) for a few genera: Chelodesmus Cook,
1895, Leptodesmus deSaussure, 1859, Odontopeltis,
Odontotropis Humbert & deSaussure, 1869,
Priodesmus Cook, 1895, Rhachodesmus Cook, 1895,
and Strongylodesmus deSaussure, 1859, without
giving a morphological diagnosis. Attems (1898) did
not adopt the family concept of Cook (1895), but
recognized 13 subfamilies in the family Polydesmidae. In that work, Attems proposed the subfamily
Leptodesminae, containing the genera Leptodesmus,
Odontotropis, Acutangulus Attems, 1899, Centrogaster
Attems, 1898, and Rhachidomorpha deSaussure,
1860. Attems (1914) and Brölemann (1916) independently elevated the Leptodesminae to family rank.
Later, Hoffman (1950) established the priority of the
family name Chelodesmidae, which pre-dates the
name Leptodesmidae.
There are taxonomic problems within the family
Chelodesmidae, as Hoffman (1982a) stated. Hoffman
(1980) divided the family Chelodesmidae into two
subfamilies, the Neotropical Chelodesminae and the
African Prepodesminae. Within the Chelodesminae,
he recognized 11 tribes. In subsequent studies,
Hoffman (1981a, b, c, 1982b, 1990a, b, 1995, 2009)
modified the tribal classification, and the number of
tribes increased to 18.
Initially, Hoffman (1980) placed the genus
Odontopeltis in the tribe Telonychopodini, a taxon
introduced by Verhoeff (1951) as Telonychopidae to
host Telonychopus meyeri Verhoeff, 1951. Hoffman
(1965a) formally proposed the tribe Telonychopodini –
with the genera Telonychopus, Catharodesmus
Silvestri, 1897, Euthydesmus Silvestri, 1902, and
Manfrediodesmus Schubart, 1943 – and subsequently
added to the same tribe the genera Odontopeltis,
Leiodesmus
Silvestri,
1897,
Brachyurodesmus
Silvestri, 1902, Eucampesmella Schubart, 1955, and
Macrocoxodesmus Schubart, 1947 (Hoffman, 1980). In
a critical re-analysis, Hoffman (2000) presented a new
synopsis for the Telonychopodini, excluding the genera
Odontopeltis, Leiodesmus (syn. Catharodesmus), and
Brachyurodesmus (not reallocated to another tribe),
placing Macrocoxodesmus and Eucampesmella into
the new tribe Macrocoxodesmini (Hoffman, 1990a),
and adding the genus Pantanalodesmus Hoffman,
2000 to the Telonychopodini. The last additions to
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
PHYLOGENY AND REVIEW OF ODONTOPELTIS
Telonychopodini are the genera Vanzolegulus Hoffman,
2002 and Dioplosternus Hoffman, 2005 (not included in
this data set). Finally, Hoffman (2005) provided a new
diagnosis for the tribe, characterized mainly by the
elongated cyphopod projecting ventrally, the large
sclerotized sternum of the male gonopod ring, with or
without a prefemoral process in the gonopod, and the
presence of rounded paranota without projections.
TAXONOMIC
HISTORY OF THE GENUS
ODONTOPELTIS
The genus Odontopeltis has a convoluted taxonomic
history with regards to the generic name as well as to
the number of included species (Hoffman, 1981a; see
Table 1). The generic name Odontopeltis was introduced by Pocock in 1894 as a replacement name for
Rhacophorus Koch, 1847, which is pre-occupied by
the frog genus Rhacophorus Kuhl & van Hasselt,
1822 (Amphibia, Anura, Rhacophoridae). According
to Pocock (1894), members of the genus are distinguished from other genera by the ‘keels of the second
segment on the same level as those of the first
and third’. Attems (1899) treated Odontopeltis as a
subgenus of Leptodesmus deSaussure, 1859, based on
the shape of paranota, and considered Odontotropis
Humbert & deSaussure, 1869 – introduced as a
subgenus of Polydesmus Latreille, 1802−03 for
Polydesmus clarazianus – as a valid genus. Later,
Attems (1938: 200, 486) reviewed the genus
Odontotropis, realising that the name was preoccupied by Odontotropis Agassiz, 1846, Mollusca and
introduced the replacement name Storthotropis.
Hoffman (1981a) revalidated the genus Odontopeltis
and synonymized with it the genera Rhacophorus
Koch, 1847, Odontotropis Humbert & deSaussure,
1869 and Storthotropis Attems, 1938.
In total, 31 South American polydesmidan species
have been associated with this genus name over time
(Table 1). Nineteen species were originally described
in Odontopeltis: nine by Silvestri (1895a, 1895b,
1897), six by Pocock (1894, 1899, 1900), three by
Attems (1898), and one species by Hoffman (1981a).
The remaining 12 species were originally placed in
the genera Polydesmus (ten species), and one species
each in Rhacophorus and Storthotropis. Twenty of the
species either described in Odontopeltis or later associated with Odontopeltis were subsequently moved to
other genera. Table 1 lists the status of each of these
species. The species Rhacophorus decoloratus Koch,
1847, was described and figured by Koch (1863), but
never again mentioned in the literature.
Hoffman (1981a) included four species in the genus
Odontopeltis, re-describing two of them: Odontopeltis
clarazianus (Humbert & deSaussure, 1869), and
Odontopeltis giganteus (Schubart, 1949), and describing the new species Odontopeltis anchisteus. The
739
fourth species, Odontopeltis conspersus (Perty, 1833),
without specimens available to him, was presented by
Hoffman (1981a) with a diagnosis based on a drawing
by Koch (1863) that lacked data on the gonopods.
Five additional species, formerly associated with
Odontopeltis, were not mentioned by Hoffman (1981a):
Odontopeltis balzanii Silvestri, 1895a, Odontopeltis
borelli Silvestri, 1895b, Rhacophorus decoloratus
Koch, 1847, Odontopeltis gracilipes (Humbert
& deSaussure, 1870), and Odontopeltis proxima
Silvestri, 1895b (see taxonomic section below).
GONOPOD
TERMINOLOGY
In all millipedes, the gonopores open on or behind the
coxae of the second pair of legs. Except for the
Penicillata, all millipedes copulate, with males using
modified appendages for sperm transfer. The
Helminthomorpha are the most species-rich and a
very diverse clade of the Diplopoda. Males of all
helminthomorph millipedes have modified legs on
the seventh and sometimes on the eighth body ring,
which are charged with sperm-containing fluid and
inserted into the female cyphopod on or behind the
coxa of the second leg pair.
Since the earliest works on Chelodesmidae up
to contemporary treatments, the nomenclature for
gonopod components has varied. Terminology has
largely followed tradition; as for other millipede
orders, primary homology hypotheses have rarely
been discussed. Latzel (1884) was the first to explicitly state the homology of helminthomorph male
gonopods to walking legs and since then the terminology of the gonopods incorporates the terminology
for leg podomeres. The most proximal podomere of the
male gonopod, identified as the coxa, shows a variety
of shapes within the Chelodesmidae, from a simple
oval shape as in most chelodesmids to large coxae as
in Macrocoxodesmus and with coxal processes as in
Manfrediodesmus. Verhoeff (1928) termed the first
gonopodal podomere coxite, whereas Shear (2000, in
the order Chordeumatida) employed the term coxite
for processes borne on the gonopodal coxa.
The cannula (Fig. 2, C), a characteristic structure of
the Polydesmida, is a process arising from the
gonopodal coxa (Fig. 2, Cx); its tapered tip inserts into
the spermatic groove (Fig. 2, SG) of the following
podomere, identified as the gonopodal prefemur or
prefemorite (Fig. 2, Pf). The functions of the cannula
(termed solenite in Hoffman, 1965b, in Oxydesmidae)
and the spermatic groove are currently unknown;
both are assumed to support the transport of sperm
(see Enghoff, 2011, for a discussion). The term ‘spermatic groove’ is widely used in the literature. Jeekel
(1982) proposed that the spermatic groove is homologous in all polydesmoid millipedes, and for that
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
Pocock, 1900: 64
Bollman, 1888: 336
Pocock, 1894: 518
Brandt, 1839: 311
Attems, 1898: 405
Karsch, 1881: 39
Peters, 1864: 622
Silvestri, 1897: 12
Attems, 1898: 100
deSaussure, 1860: 300
Silvestri, 1895b: 4
deSaussure, 1859: 323
New species
Pocock, 1894: 516
Pocock, 1894: 514
conspersus
couloni
decoloratus
discrepans
donabeja
eimeri
formosus
gayanus
giganteus
gracilipes
grantii
incises
macconnelli
magnus
mammatus
mauritii
michaelseni
morantus
mucronatus
ortonedae
polydesmoides
proxima
rubescens
sallei
salvadorii
subterraneus
tiradentes
verrucosus
vincentii
Polydesmus
Polydesmus
Rhacophorus
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Polydesmus
Storthotropis
Polydesmus
Odontopeltis
Leptodesmus
(Odontopeltis)
Odontopeltis
Rhacophorus
Odontopeltis
Polydesmus
Odontopeltis
Polydesmus
(Rhacophorus)
Polydesmus
Odontopeltis
Odontopeltis
Odontopeltis
Polydesmus
Polydesmus
(Leptodesmus)
Odontopeltis
Polydesmus
Odontopeltis
Odontopeltis
Odontopeltis
Silvestri, 1895b: 5
Gervais, 1836: 379
New species
Hoffman, 1981a: 58
Silvestri, 1895b: 4
Silvestri, 1895a: 769
Silvestri, 1895b: 3
Silvestri, 1895a: 770
Silvestri, 1895b: 4
New species
Humbert & deSaussure,
1869: 152
Perty, 1833: 210, tab
XL, fig 8.
Humbert & deSaussure,
1869: 151
Koch, 1847: 137
Silvestri, 1895b: 5
New species
Attems, 1898: 400
Pocock, 1894: 517
Gervais, 1847: 114
Schubart, 1949: 22
Humbert & deSaussure,
1870: 172
Pocock, 1899: 9
Attems, 1898: 398
aleijadinho
anchisteus
argentinea
balzanii
borellii
bovei
cameranii
xica
clarazianus
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Odontopeltis
Polydesmus
Author, year:
page number
Species
Original
genus
Pocock, 1894: 512
Schubart, 1954: 142
Listed as P. (Rhacophorus)
by Peters, 1864: 537
Pocock, 1894: 512
Attems, 1898: 402
Attems; 1898; 403
Pocock, 1894: 513
Pocock, 1894: 513
Attems, 1898: 406
Hoffman, 1981a: 60
Attems, 1898: 399
Pocock, 1894: 513
Pocock, 1909: 168
Hoffman, 1981a: 57
Silvestri, 1895b: 3
Transferred to
Odontopeltis by
Brölemann, 1909: 69
Leptodesmus
Hoffman 1982b: 256
Peréz-Asso, 1996: 189
Chamberlin, 1918: 235
Chamberlin, 1918: 236
Caraibodesmus
Antillodesmus
Loomis, 1936: 136
Schubart, 1945: 62
Attems, 1938: 117
Hoffman, 1982: 649
Silvestri, 1903: 6, syn.
of gayanus
Sandalodesmus
Amphelictogon
Quisquicia
Pseudoleptodesmus
Trichomorpha
Cordilleronomus
Trienchodesmus
Raima
Amphelictogon
Caraibodesmus
Ricodesmus
Monenchodesmus
Caraibodesmus
Hoffman, 1990b: 300
Brölemann, 1909: Catalogos
da Fauna Brazileira,
2: 69, 72
Hoffman, 1978: 551
Peréz-Asso, 1996: 195
Chamberlin, 1918: 235
Hoffman, 1999: 291
Silvestri, 1903: 8
Chamberlin, 1918: 235
Hoffman, 2000: 102
Chamberlin, 1918: 235
Silvestri, 1903: 6
Atlantodesmus
Caraibodesmus
Trienchodesmus
Fontariopsis
Synonymized with
Leptodesmus rubescens
Hoffman, 1981c: 92
Chamberlin, 1918: 232
Hoffman, 1981c: 176
Hoffman, 1981c: 92
Schubart, 1954: 122
Author of transfer
or synonymy
Stachyproctus
Amphelictogon
Strongylomorpha
Stachyproctus
Platinodesmus
Most recent transfer
or synonymy
Table 1. Species either described or placed temporarily in Odontopeltis; currently valid species of Odontopeltis in bold
Paraguay
Cuba
Brazil
Jamaica
St. Vincent and
the Grenadines
Mexico
Argentina
Brazil
Colombia
Ecuador
Chile
Guyana
Cuba
Jamaica
Puerto Rico
Chile
Jamaica
Socotra
Brazil
Unknown
Paraguay
Brazil
Brazil
Jamaica
Chile
Brazil
Brazil
Cuba
Brazil
Brazil
Brazil
Argentina
Bolivia
Paraguay
Argentina
Paraguay
Brazil
Brazil
Country
Hoffman, 1999: 270
Hoffman, 1999: 283
Hoffman, 1999: 291
Not Odontopeltis
Dalodesmidae
Dalodesmidae
Hoffman, 1999: 269
Hoffman, 1999: 269
Oxydesmidae
Not Odontopeltis
Hoffman, 1999: 269
Dalodesmidae
incertae sedis, type lost
synonym of cameranii
Neotype here
selected
Hoffman, 1999:280
Not Odontopeltis
Not Odontopeltis
Notes
740
J. P. P. PENA-BARBOSA ET AL.
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
PHYLOGENY AND REVIEW OF ODONTOPELTIS
Figure 2. Chelodesmidae gonopod terminology, Eucampesmella sp. C, cannula; Cx, coxa; Pf, prefemoral region;
PfP, prefemoral process; S, solenomere; SG, spermatic
groove (after Hoffman, 1965a, 1982b).
reason, the location of each process in relation to this
channel defines its morphological identity.
All gonopodal podomeres distal to the coxite are
termed collectively the telopodite (not to be confused
with the telopod in the Pentazonia). The proximal
component of the telopodite, usually carrying setae
(bristles), was termed ‘Schenkel’ by Attems (1894).
Later, Brölemann (1929) and Attems (1938) designated that same region as the prefemoral region
(Fig. 2, Pf), a designation still used today (e.g. see
Hoffman, 1965b: 15). However, its homology with the
prefemur of the walking legs is uncertain. Mapping
the bristle pattern of the gonopods of Polydesmida at
different ontogenetic stages, Petit (1976) proposed
that the entire telopodite in Polydesmida is homologous to the prefemur alone. This assumption implies
that in the order Polydesmida, all gonopodal branches
beyond the prefemur possibly represent evolutionary
novelties and cannot be homologized with walking leg
podomeres. Simonsen (1990), in his ‘Phylogeny of the
Polydesmidea’, accepted Petit’s hypothesis in his
phylogenetic analysis of the polydesmidan suborders,
thereby questioning Hoffman’s (1965b) and Jeekel’s
(1982) primary homology hypotheses between leg
podomeres and telopodite sections.
In many chelodesmids and other families, e.g. in
the Xystodesmidae, the telopodite carries a basal
741
Figure 3. Chelodesmid gonopod terminology comparison
and associated acropodite−podomere homology hypotheses. A, German: Verhoeff (1928, 1938), B, Brölemann
(1929), C, Attems (1894, 1938); D, Portuguese: Schubart
(1949), English: Hoffman (1965a, 1982b). Petit (1976) considered all acropodite processes as derivatives of the
prefemur. Tt, tibiotarsus.
process, called the prefemoral process (Fig. 2, PfP) by
Attems (1938). The prefemoral process sensu Attems
displays a great variety of shapes within members of
the family and its function is uncertain. However, the
homology of what has been termed the ‘prefemoral
process’ in various families of the Polydesmida is
currently unknown.
Hoffman (1965b: 15) recognized that gonopod
sections distal to the prefemur are difficult to
homologize with confidence, as some processes may be
present or absent in various chelodesmid groups, or
display widely varying forms. Frequently, the torsion
of the telopodite further obscures the identity of the
various telopodite branches. The set of distal components (Fig. 3) beyond the prefemoral region are
called the acropodite (Ribaut, 1920; Verhoeff, 1928;
Hoffman, 1982a). Schubart (1949) and Hoffman
(1965b) identified the same region as the femoral
region, characterized by the absence of bristles. In the
Chelodesmidae, the number of processes present in
the acropodite can vary from one to three. The main
process is called the solenomere, defined as the
branch or process that carries the seminal (spermatic)
groove. It has also been called the Rinnenast
(German, e.g. Attems, 1938) or Solänomerit (Verhoeff,
1938).
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
742
J. P. P. PENA-BARBOSA ET AL.
apical processes, the solenomere (S), process A
(Fig 4, PrA), and process B (Fig. 4, PrB). The apical
processes have been named differently by various
authors. Schubart (1949) identified the three apical
processes as the femoral process (proximal, PrB),
tibiotarsus (median, PrA), and solenomere (external,
S, Fig. 4A, B), implying homology with telopodite
processes present in other Chelodesmidae. Hoffman
(1981a), as Brölemann (1900) before, avoided terminology that could imply such homology and called
these collectively ‘apical processes’; Brölemann
labelled the branches alphabetically, consistent with
our terminology employed here.
Figure 4. Gonopod structures of Odontopeltis, left
gonopod. A, Odontopeltis conspersus, mesal view. B,
Odontopeltis anchisteus, ventral view. AB, acropodite bristles; AF, acropodite folds; BPPf, basal projection of
prefemoral region; C, cannula; Cx, coxa of gonopod; FR,
femoral region; Pf, prefemoral region; PrA, process A of
the acropodite; PrB, process B of the acropodite; S,
solenomere; SG, spermatic groove; SP, spiniform projection on gonopod coxa. Scale bar = 0.5 mm.
Attems (1938) proposed that the solenomere must
be considered a new process, nonhomologous to
any other structure of the walking legs. By contrast,
Hoffman (1965b) assumed that the solenomere is
homologous to the distal end of the walking leg, the
tarsus, because the spermatic groove would logically
run to the tip of a modified walking leg; therefore, the
distal end of the spermatic groove most likely identifies the tarsus tip of the walking leg. Later, Hoffman
(1990b) discussed several possible homology hypotheses for the solenomere, sometimes associated with
the tarsus and/or the tibio-tarsus. Jeekel considered
the presence of the tibiotarsal process in addition
to the solenomere and other prefemoral or femoral
processes (Jeekel, 1968: 22 for Paradoxosomatidae)
and thus did not assume homology between the
solenomere and tibiotarsus.
DESCRIPTION OF THE MALE GONOPOD
OF ODONTOPELTIS
The gonopods in Odontopeltis males (Fig. 4) consist of
a single cylindrical branch; a basal prefemoral process
is absent. The gonopod coxae may possess a spiniform
process (Fig. 4, SP). The spermatic groove runs on the
mesal side of the acropodite. The telopodite shaft can
be divided into two regions: the prefemoral (Pf) and
the femoral region (FR). The boundary is demarcated
by two long bristles (AB) in a transverse or inclined
line. The mostly setose Pf possesses an angular basal
projection (BPPf). The ectal side of the prefemoral
region may display several folds (AC) in the central
section of the telopodite shaft. The FR carries three
MATERIAL AND METHODS
MORPHOLOGICAL EXAMINATIONS
Morphological observations and illustrations were
undertaken using a Leica MZ12 stereomicroscope
with a camera lucida. Digital scanning electron
microscopy (SEM) photographs were taken on a
Zeiss DSM 940 scanning electron microscope at
the Laboratório de Microscopia do Instituto de
Biociências, Universidade de São Paulo. All measurements are in millimetres. Gonopod terminology
follows Brölemann (1900) and Hoffman (1981a),
whereas body terminology follows Attems (1898) and
Brölemann (1900). The paranota carry a variable,
often species-specific number of tooth-like lateral projections, which are denoted in a schema following
Schubart (1949): figures indicate the total number of
tooth-like projections at the paranotum on each body
ring, e.g. ‘0’ indicates ‘no tooth-like projection’.
CLADISTIC
ANALYSIS
The data matrix comprises 47 characters and 15
terminals (Table 2), edited in MESQUITE 2.0
(Maddison & Maddison, 2010), in which nonapplicable and unknown data are presented as ‘-’ and
‘?’, respectively. The 15 terminals used on this analysis comprise all four species currently allocated
to Odontopeltis (Hoffman, 1981a), as well as the
four new species described here. The existing tribal
structure, in which the many chelodesmid genera
are placed, is currently not based on putative
apomorphies. To fully investigate the phylogenetic
affinities of Odontopeltis would require a complete
cladistic analysis of the majority of the chelodesmid
genera. This task is beyond the scope of this study.
Therefore, we selected six species as outgroup terminals, belonging to genera that had previously been
placed close to Odontopeltis by Hoffman (1980;
see Table 3): Eucampesmella, Macrocoxodesmus,
Manfrediodesmus, Pantanalodesmus, Rondonaria,
and Telonychopus. The species Leiodesmus major
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
743
Table 2. Graphical representation of character support for clades identified in Figure 5: cell shading denotes character
state: open cell, state 0; grey cell, state 1; black cell, state 2; NA, not applicable; ?, unknown. Clades named as in Figure 5
4
18
38
7
40
33
35
14
27
3
15
25
39
5
6
47
8
24
32
9
11
17
19
20
31
46
23
26
12
16
22
30
10
13
45
21
43
29
28
1
2
34
36
37
41
42
44
o. ale ijadinho
O . tirade nte s
o. c hic a
o. donabe ja
o. c laraz ianus
o. anc his te us
o.c ons pe rs us
O . gigante us
R ondanaria
Mac roc ox ode s m u
C la d e G
C la d e F
O d o n t o p e lti s
C la d e A
C la d e C
C la d e B C la d e D C la d e E
Manfre diode s m us
T e lony c hopus
Pantanalode s m us
E uc am pe s m e lla
L e iode s m us
c ha ra c te r
C la d e H
?
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
?
NA
?
?
?
?
?
?
NA
?
?
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
?
?
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
?
?
NA
?
?
NA
NA
NA
NA
NA
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
744
J. P. P. PENA-BARBOSA ET AL.
Table 3. List of outgroups used in the cladistic analysis
Family
Tribe
Species
Depository
Chelodesmidae
Chelodesmidae
Chelodesmidae
Chelodesmidae
Chelodesmidae
Chelodesmidae
Chelodesmidae
no tribe assigned, Hoffman, 2000: 3
Macrocoxodesmini
Macrocoxodesmini
Telonychopodini
Telonychopodini
Telonychopodini
no tribe assigned, Hoffman, 2006
Leiodesmus major
Eucampesmella ferrii
Macrocoxodesmus marcusi
Telonychopus klossae
Manfrediodesmus passarelii
Pantanalodesmus marinezae
Rondonaria schubarti
MRSN
MZSP
MZSP
MZSP
MZSP
FMNH
MZSP
Figure 5. Implied weighting tree, showing the relationship of the genus Odontopeltis and the outgroup taxa. White
(homoplasious) and black (nonhomoplasious) circles represent character transformations. Number above the circle
indicates character number; number below the circle indicates character state. Bremer values: numbers in parentheses.
Capital letters indicate clades discussed in the text.
Silvestri, 1897 was used to root the tree based on
taxonomic position as a genus once placed in the
Telonychopodini (Hoffman, 1980) and subsequently
removed (nontribal position, Hoffman, 1981a).
After construction of the data matrix, the analysis
was performed in TNT software (Goloboff, Farris &
Nixon, 2003–2004), using parsimony (Farris, 1983)
and the implied weighting technique (Goloboff, 1993)
to search for the most parsimonious tree, with concavity value (k) equal to 2.012 (according to Mirande,
2009). Branch support is expressed by Bremer support
values (Bremer, 1994) and presented as a number
between parentheses at each node of Figure 5.
ABBREVIATIONS
Morphological abbreviations
A, acropodite; AB, acropodite macrobristles; AF,
acropodite folds; BPPf, basal projection of prefemoral
region; C, cannula; CS, central support of gonopod
aperture; CT, coxal tip; Cx, coxae; FR, femoral region;
IPCx, internal projection of coxae; LF, lateral folds of
gonopod aperture; MPS, median process of solenomere;
Oz, ozopore; Pf, prefemoral region; PfP, prefemoral
process; PrA, process A; PrB, process B; PB, posterior
border of collum; PE, posterior edge of the gonopod
aperture; S, solenomere; SB, solenomere base; SG,
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
745
spermatic groove; SP, spiniform process of the gonopod
coxa; T, telopodite; Tt, tibiotarsus; TO, Tömösváry
organ.
Museum abbreviations (curators or
collection managers)
BMNH, The Natural History Museum, London, UK
(J. Beccaloni); IBSP, Instituto Butantan, São Paulo,
Brazil (D. M. Barros Battesti); MHNG, Muséum
d’Histoire Naturalle, Geneva, Switzerland (P.
Schwendinger); MNHN, Muséum National d’Histoire
Naturelle, Paris, France (J. J. Geoffroy); MNRJ,
Museu Nacional do Rio de Janeiro, Rio de Janeiro,
Brazil (A. B. Kury); MRSN, Museo Regionale di
Scienze Naturali, Torino, Italy (A. Chiarle); MZSP,
Museu de Zoologia de São Paulo, São Paulo, Brazil
(R. Pinto da Rocha); UFMG, Universidade Federal de
Minas Gerais, Belo Horizonte, Brazil (A. J. Santos);
ZSM,
Zoologische
Staatssammlung
München,
Germany (Dr Roland Melzer); ZMB, Zoologisches
Museum Berlin, Germany (Dr Jason Dunlop); ZIN,
Zoological Institute, St. Petersburg, Russia (Dr Victor
A. Krivokhatsky); ZMUZ, Zoologisches Museum der
Universität Zürich, Switzerland (Dr Paul Ward).
RESULTS AND DISCUSSION
CHARACTERS
A list of the characters used in this analysis is given
below with their respective length (L) plus consistency
(CI) and retention (RI) indexes in this data set. As all
characters listed here are used for the first time in a
phylogenetic analysis within Chelodesmidae, character state changes are discussed. Character state
changes are mapped on the resulting cladogram
(Fig. 5).
HEAD
1. Row #1 of bristles on head (Fig. 6): present (0);
absent (1). L = 3; CI = 0.33; RI = 0.33.
Some Polydesmida show series of rows of bristles
on the dorsal surface of the head. Row 1 is located
between the antennae. In this analysis there is a
secondary loss of row 1 in Rondonaria schubarti,
reappearing in Odontopeltis tiradentes.
2. Number of bristle rows anterior of row #1: two
rows (0); three rows (1). L = 2; CI = 0.50; RI = 0.
All Chelodesmidae show a series of bristles right
below row #1 (Fig. 6). A third row appears independently in Manfrediodesmus passarelii and in
Macrocoxodesmus marcusi.
3. Shape of Tömösváry organ (Figs 7, 8): circular (0);
suboval (1). L = 1; CI = 1; RI = 1.
Figures 6–11. Head characters of Odontopeltis. Figure 6.
Dorsal face of head of Odontopeltis giganteus, showing the
rows of bristles (documenting characters 1 and 2 of the
phylogenetic analysis). I, first row between antennal
sockets; II, second row; III, third row. Figures 7, 8.
Chelodesmid head, schematic, lateral view, showing
shape of the Tömösváry organ (TO, character 3).
Figure 7. Suboval. Figure 8. Circular. Figures 9–11. Last
antennomere of chelodesmids, showing the four apical
sense cones (character 4). Figure 9. Double invagination.
Figure 10. Triple invagination. Figure 11. Quadruple
invagination.
The ‘suboval’ state appears to be an exclusive
apomorphy of Odontopeltis (Fig. 7).
ANTENNAE
4. Number of invaginations between the four apical
sense cones on the antenna (Figs 9–11): two (0);
three (1); four (2). L = 4; CI = 0.50; RI = 0.33.
Almost all taxa used in this analysis share a pair
of invaginations between the apical sense cones
on the antennae. A third invagination occurs
independently in Rondonaria schubarti and in
Clade E (Fig. 5): Odontopeltis tiradentes + Odontopeltis aleijadinho. A fourth invagination
appears independently in Telonychopus klossae
and in Odontopeltis giganteus.
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J. P. P. PENA-BARBOSA ET AL.
8.
9.
10.
Figures 12–18. Body ring characters. Figures 12, 13.
Collum shape of chelodesmids, showing the posterior edge
(PB) (character 5). Figure 12. Straight. Figure 13. Arched.
Figures 14–16. Tip of seventh paranota in Odontopeltis
species, showing the shape of the yellow spot (dashed line,
characters 7, 12). Figure 14. Subrectangular, Odontopeltis
anchisteus. Figure 15. Subtriangular, Odontopeltis donabeja. Figure 16. Semicircular, Odontopeltis clarazianus.
Figures 17, 18. Ventral projections at the fifth body ring
(character 8). Figure 17. Four projections. Figure 18. Two
projections.
11.
BODY
5. Posterior edge of collum (Figs 12, 13): arched (0);
straight (1). L = 2; CI = 0.50; RI = 0.80.
The ‘arched’ state is present in a great number of Chelodesmidae species, whereas, in this
analysis, the ‘straight’ state appears independently in Eucampesmella ferrii and Odontopeltis
(Fig. 5).
6. Coloration of edge of paranota (Fig. 1): absent (0);
present (1). L = 2; CI = 0.50; RI = 0.80.
This characteristic feature (Fig. 1) is present in
the genera Odontopeltis and Eucampesmella. The
analysis revealed an independent origin (Fig. 5).
7. Shape of the colour spot on the seventh paranotum
(Figs 14–16): subrectangular (0); subtriangular
(1); semicircular (2). L = 3; CI = 0.66; RI = 0.
The plesiomorphic ‘subrectangular’ state is
present in E. ferrii and is also observed in some
Odontopeltis species. The subtriangular state is
homoplastic, occurring in Odontopeltis aleijadinho
and in Odontopeltis anchisteus; it changes to the
12.
13.
autapomorphic semicircular state in Odontopeltis
clarazianus.
Number of ventral projections on the fifth body
ring (Figs 17, 18): two (0); four (1). L = 2;
CI = 0.50; RI = 0.75.
Males of Chelodesmidae bear either one or two
pairs of projections on the sternites of the
fifth body ring, between the legs. One pair of
ventral projections on the fifth body ring occurs
widely in the family. An additional pair, the
apomorphic condition, occurs independently in
Pantanalodesmus marinezae and in the genus
Odontopeltis.
Pair of ventral projections on sixth body ring:
absent (0); present (1). L = 2; CI = 0.50; RI = 0.75.
Clade F (Rondonaria schubarti + Odontopeltis)
shows a pair of ventral projections on the
sixth body ring, which is also present in
Pantanalodesmus marinezae.
Pair of ventral projections on the seventh body
ring: absent (0); present (1). L = 2; CI = 0.50;
RI = 0.66.
A few species of Chelodesmidae carry a pair of
projections right below the gonopod aperture. The
presence of these projections is homoplastic,
appearing independently in Leiodesmus major
and in the clade Telonychopodini.
Position of the ozopore on the edge of paranota
(Figs 19–21): anterior (0); medial (1); posterior
(2). C = 4; CI = 0.50; RI = 0.33.
The position of the ozopore on the edge of
paranota varies among members of the Chelodesmidae. The ‘anterior position’ state is present
in the sister-group Telonychopus klossae + Manfrediodesmus passarelii and in Macrocoxodesmus
marcusi. The ‘medial position’ state is found in
Eucampesmella ferrii and in Clade F: Rondonaria
schubarti + Odontopeltis. The ‘posterior position’
appears in Leiodesmus major and Pantanalodesmus marinezae independently.
Projections on the edge of paranota (Figs 14–16):
absent (0); present (1). L = 1; CI = 1; RI = 1.
Projections on the edge of paranota are not
common amongst the Chelodesmidae, but are
more frequently observed in the Platyrhacidae. In
this analysis, the presence of projections on the
paranota edge appears as a synapomorphy for
Clade G: Macrocoxodesmus marcusi + Rondonaria
schubarti + Odontopeltis.
Alignment of paranota in posterior view (Figs 22–
24): straight (0); curved ventrally (1); curved dorsally (2). L = 2; CI = 1; RI = 1.
The state ‘straight paranota’ is plesiomorphic
in this analysis. Two independent state changes
occur within this data set: in the Telonychopodini
the paranota curve ventrally, whereas in
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
747
data set, the presence of a dorsal lobe on
the prefemur of the fifth leg represents a
synapomorphy of Clade G: Macrocoxodesmus
marcusi + Rondonaria schubarti + Odontopeltis.
17. Ventral bristles of legs (Figs 27, 28): heavy and
large (0); thin and small (1). L = 2; CI = 0.50;
RI = 0.75.
Here, the ‘heavy and large’ state is plesiomorphic,
with two independent changes to the derived state
‘thin and small’: in Eucampesmella ferrii and in
Clade F: Rondonaria schubarti + Odontopeltis.
MALE GONOPORE
Figures 19–24. Body ring characters. Figures 19–21.
Ozopore position (Oz) on the paranota (character 11).
Figure 19. Anterior. Figure 20. Median. Figure 21. Posterior. Figures 22–24. Alignment of paranota in Polydesmida
(character 13). Figure 22. Raised, Macrocoxodesmus
marcusi; Figure 23. Straight, Odontopeltis giganteus.
Figure 24. Lowered, Telonychopus klossae.
Macrocoxodesmus marcusi the paranota curve
dorsally.
14. Number of projections at the edge of seventh
paranotum (Figs 14–16): one (0); two (1); three or
more (2). L = 4; CI = 0.50; RI = 0.50.
‘One projection’ is the plesiomorphic state in
this analysis; there is a state change in
Macrocoxodesmus marcusi which features ‘two
projections’. The state ‘three or more projections’
represent a synapomorphy for Clade A within
Odontopeltis, with two independent reversals to
‘two projections’: one in Odontopeltis anchisteus
and another in Clade D: Odontopeltis
donabeja + Odontopeltis xica.
15. Ozopore rim (Figs 25, 26): raised (0); not raised
(1). L = 1; CI = 1; RI = 1.
The ozopore rim in the Chelodesmidae usually
shows a raised edge. The loss of the raised edge is
a synapomorphy of Odontopeltis.
16. Dorsal lobe on prefemur of fifth leg (Figs 31, 32):
present (0); absent (1). L = 1; CI = 1; RI = 1.
The prefemora of legs in Chelodesmidae are
usually straight, without a dorsal lobe. In this
18. Male genital papilla dimensions: height equal to
width of base (0); base twice as wide as total
height (1); height larger than base width (2).
L = 5; CI = 0.40; RI = 0.50.
The state 0 ‘height equal to width of base’ appears
three times in this data set: in Leiodesmus
major, in Pantanalodesmus marinezae, and in
Odontopeltis. However, within Odontopeltis there
are two additional state changes: Odontopeltis
clarazianus displays an autapomorphic state, the
height of the genital papilla being larger than the
width of the base (state 2). State 1 ‘base twice as
wide as total height’ is present in Eucampesmella
ferrii, Telonychopus klossae, Manfrediodesmus
passarelii, Macrocoxodesmus marcusi, Rondonaria schubarti, and Clade E (Odontopeltis
tiradentes + Odontopeltis aleijadinho).
19. Shape of male genital papilla (Figs 29, 30): rectangular (0); conical (1). L = 1; CI = 1; RI = 1.
The genital papilla shows two basic shapes in
the Chelodesmidae. The conical shape (state 1)
is a synapomorphy for Clade F (Rondonaria
schubarti + Odontopeltis). The rectangular shape
(state 0) is a plesiomorphy shared by other outgroup species.
GONOPODS
20. Lateral folds at the posterior border of the
gonopod aperture (Fig. 38, LF): present (0);
absent (1). L = 1; CI = 1; RI = 1.
The aperture of the gonopod is oval in Polydesmida, with family level variations (Hoffman,
1980). In some cases within Chelodesmidae,
the aperture possesses a few lateral folds.
The presence of lateral folds is a plesiomorphy
present in Leiodesmus major, E. ferrii, members
of the Telonychopodini, and Macrocoxodesmus
marcusi. Their absence is a synapomorphy of
Clade F (Rondonaria schubarti + Odontopeltis).
21. Number of lateral fold at the posterior border of
gonopod aperture: three or fewer (0); more than
five (1). L = 1; CI = 1; RI = 1.
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J. P. P. PENA-BARBOSA ET AL.
Figures 25–28. Somatic characters. Figures 25, 26. Scanning electron microscopy (SEM) images of paranota, showing
the two states of the ozopore rim found in Chelodesmidae (character 14). Figure 25. Raised, Manfrediodesmus passarelii.
Scale bar = 0.5 mm. Figure 26. Flat, Odontopeltis donabeja. Scale bar = 0.5 mm. Figures 27, 28. SEM images of different
forms of ventral leg bristles (character 17). Figure 27. Heavy and large, Manfrediodesmus passarelii. Scale bar = 0.5 mm.
Figure 28. Thin and small, Odontopeltis donabeja. Scale bar = 0.3 mm.
The state (1) ‘more than five’ folds appears as
a synapomorphy in Clade H (Eucampesmella
ferrii + Telonychopodini).
22. Central support of gonopod aperture (Figs 38, 39,
CS): sclerotized (0); membranous (1). L = 1;
CI = 1; RI = 1.
In Polydesmida, the gonopod aperture features
a region for the support of the gonopod, which
may be membranous or sclerotized. The state
‘membranous’ is a synapomorphy for the
clade Macrocoxodesmus marcusi + Rondonaria
schubarti + Odontopeltis; and ‘sclerotized’ appears
as a plesiomorphy in Leiodesmus major,
Eucampesmella ferrii, and Telonychopodini.
23. Posterior margin of gonopod aperture (Figs 38–
40, PE): deeply excavated, surpassing half of the
coxae of ninth pair of legs (0); excavation reaching half of the coxae of ninth pair of legs (1);
excavation shallow, not reaching the coxae of
ninth pair of legs (2). L = 2; CI = 1; RI = 1.
As discussed by Simonsen (1990), the suborder Chelodesmidea is characterized by large,
protuberant gonopod coxae, located almost completely outside the gonopod aperture. In the
Chelodesmidae there is great variation in the
shape of the coxae of the gonopods, embedded in
a gonopod aperture that often features accommodations for these protuberant coxae. Thus, species
that possess large coxae tend to have a deeply
excavated posterior margin of the gonopod aperture. Here, state 0 ‘deeply excavated, surpassing
half of the coxae of ninth pair of legs’ represents
a plesiomorphy present in Leiodesmus major,
E. ferrii, Pantanalodesmus marinezae, and
Macrocoxodesmus marcusi, coincidentally the
species with the largest gonopod coxae. Within
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
749
Figures 29, 30. Scanning electron microscopy images of the two alternate shapes of the genital papilla found in males
in this data set (character 19). Figure 29. Conical shape, Odontopeltis tiradentes. Scale bar = 0.1 mm. Figure 30.
Rectangular shape, Manfrediodesmus passarelii. Scale bar = 0.1 mm.
the Telonychopodini there is a state change
to state 1 ‘excavation reaching half of the
coxae of ninth pair of legs’ characterizing the
clade Telonychopus klossae + Manfrediodesmus
passarelii. The state ‘excavation shallow, not
reaching the coxae of ninth pair of legs’ appears
as a synapomorphy for clade F (Rondonaria
schubarti + Odontopeltis).
24. Prefemoral process (Fig. 2): present (0); absent
(1). L = 2; CI = 0.50; RI = 0.66.
As discussed in the section on gonopod morphology and terminology, many members of
the Chelodesmidae possess large prefemoral
processes in the gonopods, representing the
plesiomorphic condition within the family. In
this data set, the absence of a prefemoral process
occurs twice independently, in the Telonychopodini and in Odontopeltis.
25. Ectal folds on the telopodite (Fig. 4): absent (0);
present (1). L = 1; CI = 1; RI = 1.
The presence of folds (AF) on the ectal side of the
telopodite shaft is an exclusive synapomorphy for
Odontopeltis.
26. Position of solenomere in relation to other
acropodite processes (Fig. 2): medial, surrounded
by basal processes (0); distal, exposed (1). L = 3;
CI = 0.33; RI = 0.33.
The ‘distal’ state appears, independently,
two times in this analysis: an apomorphy in
the Telonychopodini, with a reversal to the
27.
28.
29.
30.
‘medial’ state in Telonychopus klossae. Clade F
(Rondonaria schubarti + Odontopeltis) is characterized by the apomorphic state, the exposed
solenomere.
Spiniform process on gonopod coxa (Figs 33, 34):
absent (0); present (1). L = 2; CI = 0.50; RI = 0.83.
The coxal spiniform apophysis is a homoplastic
character, which evolved independently in
Telonychopus klossae and in Clade A within
Odontopeltis.
Size of spiniform process on gonopod coxa
(Figs 33, 34): short – half of the length of ‘long’
state (0); long (1). L = 3; CI = 0.33; RI = 0.
The ‘long’ spiniform coxal process appears
three times independently within this data
set: in Clade H, Odontopeltis clarazianus, and
Odontopeltis donabeja.
Solenomere shape (Figs 35–37): suboval (0);
falciform (1); sinuous (2). L = 4; CI = 0.50; RI = 0.
The ‘falciform’ state appears as a synapomorphy
of all species, except Leiodesmus major, with
three independent changes, in Eucampesmella
ferrii to ‘suboval’, and two independent state
changes to ‘sinuous’ in Pantanalodesmus
marinezae and Macrocoxodesmus marcusi.
Spines on solenomere (Figs 41, 42): covering the
solenomere (0); within the spermatic groove (1).
L = 2; CI = 0.50; RI = 0.50.
Spines covering the solenomere appear to be a
plesiomorphy, with two character state changes to
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J. P. P. PENA-BARBOSA ET AL.
34.
35.
Figures 31–37. Leg and gonopod characters. Figures 31,
32. States of the prefemur in Chelodesmidae (character
16). Figure 31. Dorsally globose. Figure 32. Dorsally
straight. Figures 33, 34. Spiniform process (SP) on the
coxa (Cx) of the gonopod (characters 27, 28). Figure 33.
Short – half of the length of the long state. Figure 34.
Long. Figures 35–37. Femoral region of the gonopod
showing three forms of the solenomere (S) present in
this analysis (character 29). Figure 35. Suboval,
Leiodesmus major. Figure 36. Falciform, Telonychopus
klossae. Figure 37. Sinuous, Pantanalodesmus marinezae.
C, canula; Pf, prefemur.
the derived condition: in Telonychopodini and in
Clade F (Rondonaria schubarti + Odontopeltis).
31. Solenomere thickness: thick (0); flat (1). L = 1;
CI = 1; RI = 1.
The ‘flat’ state is a synapomorphy for Clade F
(Rondonaria schubarti + Odontopeltis).
32. Median projection of the solenomere (Figs 43–45):
absent (0); present (1). C = 4; CI = 0.25; RI = 0.50.
The median projection on the solenomere evolved
two times independently in Telonychopus klossae
and as a synapomorphy for Odontopeltis, with
two independent reversals in Odontopeltis
anchisteus and in Odontopeltis tiradentes.
33. Shape of median projection on solenomere:
rounded (0); pointed (1). L = 1; CI = 1; RI = 1.
Within the genus Odontopeltis the ‘pointed shape’
appears as a synapomorphy of Clade C, present
36.
37.
38.
in (Odontopeltis donabeja; Odontopeltis xica) and
Odontopeltis aleijadinho; Odontopeltis tiradentes
does not display any kind of median projection on
the solenomere.
Size relationship between the solenomere and
median projection (Figs 43–45): solenomere larger
than median projection (0); solenomere and
median projection equal in size (1); solenomere
smaller than median projection (2). L = 3;
CI = 0.66; RI = 0.
This character is rather homoplastic in this
data set. State 1 ‘solenomere and median projection equal in size’ has a homoplastic distribution, occurring independently in Odontopeltis
conspersus and Odontopeltis aleijadinho. Odontopeltis donabeja shows the autapomorphic state 2:
‘solenomere smaller than median projection’.
Odontopeltis giganteus, Odontopeltis clarazianus,
and Odontopeltis xica share the plesiomorphic state 0 ‘solenomere larger than median
projection’.
Extension of solenomere base (Figs 46–48): not
reaching half of the width of process A (0); reaching half of the width of process A (1); beyond half
of process A (2). L = 2; CI = 1; RI = 1.
The plesiomorphic state ‘not reaching half of the
width of process A’ is present in all outgroup taxa
and also in Odontopeltis giganteus, Odontopeltis conspersus, Odontopeltis anchisteus, and
Odontopeltis clarazianus. Character state change
to state 1 ‘reaching half of the width of process A’
appears as a synapomorphy of Clade C, with an
additional change to state 2 ‘beyond half of
process A’ as an autapomorphy in Odontopeltis
aleijadinho.
Hump on process A, in ventral view (Figs 49, 50):
absent (0); present (1). L = 2; CI = 0.50; RI = 0.50.
In this data set, the presence of a hump on
process A is an independent apomorphy for
Clade B (Odontopeltis anchisteus + Odontopeltis clarazianus) and an autapomorphy for
Odontopeltis donabeja.
Fold on process A, mesal view: absent (0);
present (1) (Fig. 62, arrow). L = 3; CI = 0.33;
RI = 0.33.
The presence of a fold on process A appears
independently twice as an apomorphic character
state change in Odontopeltis aleijadinho and in
Odontopeltis xica.
Direction of process B, mesal view (Fig. 4A): posterior (0); ectally (1); dorsal (2). L = 2; CI = 1;
RI = 1.
State 1 is an autapomorphy for Odontopeltis
giganteus (process B pointing ectally, Fig. 68).
Clade A is characterized by the synapomorphic
character state change to state 2 (e.g. Fig. 65),
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
751
Figures 38–42. Gonopod characters. Figures 38–40. Sternal gonopod opening, showing the lateral folds (LF), the central
support (CS), and the posterior excavation (PE) (characters 20, 22). Figure 38. Presence of lateral folds, central support
membranous and posterior margin deeply excavated, surpassing half of the coxae (Cx) of ninth pair of legs (Character 23),
Macrocoxodesmus marcusi. Scale bar = 1.0 mm. Figure 39. Sternal gonopod opening showing the central support
sclerotized and posterior excavation reaching half of the coxae of ninth pair of legs (Character 23), Telonychopus klossae.
Scale bar = 1.0 mm. Figure 40. Absence of lateral folds and posterior edge not reaching half of the coxa, Odontopeltis
giganteus. Figures 41, 42. Spines on solenomeres (character 30 in this analysis). Scale bar = 1.0 mm. Figure 41. Spines
within spermatic groove, Odontopeltis donabeja. Figure 42. Spines inside and outside spermatic groove, covering the
solenomere, Leiodesmus major. Scale bar = 20 μm.
with a reversal to the plesiomorphic condition in
Clade D (e.g. Fig. 77).
39. Pair of macrobristles on the telopodite shaft
(Fig. 4A, B): absent (0); present (1). L = 1; CI = 1;
RI = 1.
The presence of macrobristles is a synapomorphy
for the genus Odontopeltis.
40. Distance between the two macrobristles: less than
half the length of each macrobristle (0); more
than half the length of each macrobristle (1).
L = 1; CI = 1; RI = 1.
The state ‘less than half the length of each
macrobristle’ is a plesiomorphy within Odontopeltis. The opposite state is a synapomorphy for
Clade B.
41. Position of macrobristles: inclined (0); on same
plane (1). L = 3; CI = 0.33; RI = 0.
The ‘inclined’ (Fig. 63) state appears as a
plesiomorphy within Odontopeltis, with three
independent state changes in Odontopeltis
conspersus, Odontopeltis xica, and Odontopeltis
aleijadinho.
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J. P. P. PENA-BARBOSA ET AL.
Figures 43–50. Gonopod characters in Odontopeltis.
Figures 43–45. Solenomere (S) and median projection
(MPS) in Odontopeltis, ventral view (characters 32, 34).
Figure 43. Solenomere tip smaller than MPS of
solenomere. Figure 44. Solenomere tip and median projection equal in size. Figure 45. Solenomere tip larger than
the median projection. Figures 46–48. Extension of
solenomere basis (BS) (character 35). Figure 46. Not
reaching half of the width of process A (PrA). Figure 47.
Reaching half of the width of PrA. Figure 48. Reaching
beyond half of the width of PrA. Figures 49, 50. Hump on
PrA, ventral view (character 36). Figure 49. PrA without
apical hump. Figure 50. PrA with apical hump.
42. Trajectory of spermatic groove to the solenomere
(Figs 55, 56): straight (0); curved (1). L = 3;
CI = 0.33; RI = 0.33.
The trajectory of the spermatic groove can be
used to define homologous sections within the
gonopod structures (Jeekel, 1982). The spermatic
groove trajectory may also indicate torsion of
some part of the gonopod (Hoffman, 1990b).
Applied to the current analysis, the straight
trajectory of the spermatic groove in the
clade Eucampesmella ferrii + Telonychopodini
and, independently in Odontopeltis indicates
nontorsion of the gonopod, whereas the
curved trajectory of the spermatic groove as in
Figures 51–54. Gonopod and cyphopod characters.
Figures 51, 52. Schema of gonopod coxa in Chelodesmidae,
ectal view (character 43). Figure 51. No projection.
Figure 52. Projection. Figures 53, 54. Cyphopod structure,
posterior aspect of cyphopod in ventral view (character 46).
Figure 53. Odontopeltis giganteus, short state of cyphopod.
Figure 54. Eucampesmella ferrii, long state of cyphopod.
Manfrediodesmus passarelii may indicate torsion
of the gonopod.
43. Tip of gonopod coxae in ectal view (Figs 51, 52):
projected (0); nonprojected (1). L = 1; CI = 1;
RI = 1.
The ‘nonprojected’ state appears as a synapomorphy for the clade Eucampesmella
ferrii + Telonychopodini. All other species in this
analysis show the ‘projected’ state, with varying
degrees of length.
44. Shape of the tip of coxae tip in ectal view (Figs 51,
52): oval (0); pointed (1). L = 2; CI = 0.50; RI = 0.
The ‘pointed’ shape appears as a synapomorphy
for all species in this analysis, except Leiodesmus
major. In Clade F (Rondonaria schubarti +
Odontopeltis) there is a reversal to the ‘oval’
character state.
45. Projection of gonopod coxae, left gonopod, mesal
view: absent (0) (Fig. 56); present (1) (Fig. 55,
arrow IPCx). L = 1; CI = 1; RI = 1.
The presence of a mesal projection on the
gonopod’s coxae is a synapomorphy of the
Telonychopodini.
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753
Figures 55–58. Gonopod and body characters (scanning electron microscopy). Figures 55, 56. Left gonopods, mesal view
(characters 42, 43, 45). Figure 55. Manfrediodesmus passarelii, with projection (IPCx, internal projection of coxa: arrow)
and curved spermatic groove. Scale bar = 2.0 mm. Figure 56. Odontopeltis giganteus, without projection and straight
spermatic groove. Scale bar = 1.0 mm. Figures 57, 58. Odontopeltis donabeja. Figure 57. Tubercles on paranota, dorsal
region. Scale bar = 100 μm. Figure 58. Small spines and bristles with median projection on penultimate and ultimate
antennomeres. Scale bar = 100 μm.
CYPHOPODS
46. Size of apical projection of cyphopod (Figs 53, 54):
long (0); short (1). L = 1; CI = 1; RI = 1.
The ‘long’ state appears as a plesiomorphy,
whereas the loss of a long projection supports
Clade G (Macrocoxodesmus marcusi + Rondonaria schubarti + Odontopeltis).
47. Direction of the cyphopod aperture: ectal (0); posterior (1). L = 2; CI = 0.50; RI = 0.75.
The ‘ectal’ state appears as a plesiomorphy, with
two independent state changes to ‘posterior’, in
E. ferrii and in the genus Odontopeltis.
The cladistic analysis of 15 taxa and 47 characters,
with implied weighting, resulted in a single tree
with 99 steps (CI = 58; RI = 73). The topology sug-
gests that the tribe Telonychopodini is monophyletic,
supported by two exclusive synapomorphies: ventrally
curved paranota (character 13) and presence of a
projection on the gonopodal coxa (character 45).
Golovatch & Hoffman (2004), citing the ‘obvious’
relationship between Eucampesmella (formerly in
the Telonychopodini) and Macrocoxodesmus, also
discussed in Hoffman (1990a: 170), transferred
Eucampesmella to the tribe Macrocoxodesmini. The
topology resulting from this analysis suggests that
Eucampesmella is a sister-group of Telonychopodini
(Clade H), sustained by character 21, state 1 (more
than five lateral folds at the gonopod aperture) and
character 43, state 1 (absence of coxal tip projection).
Indeed, this latter character is listed as a diagnostic
character for Macrocoxodesmini by Hoffman (1990a).
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J. P. P. PENA-BARBOSA ET AL.
The Telonychopodini (Clade H)are sister to Clade G.
The genus Macrocoxodesmus is sister to Clade F,
consisting of Rondonaria + Odontopeltis, supported
by character 12.1 (the presence of projections on the
paranota edge), character 16.1 (prefemur of fifth legs
without dorsal lobe), character 22.1 (central support
of gonopods opening membranous), and possibly
character 46.1 (short apical cyphopod projection in
females, unknown for Macrocoxodesmus).
Clade F, encompassing the genus Rondonaria and
its sister-group, the monophyletic genus Odontopeltis,
is supported by four synapomorphies: character 19.1
(genital papilla conical), character 20.1 (lateral folds
on posterior border of gonopod opening absent), character 23.1 (concave posterior border of gonopod
opening not reaching the coxae of the ninth leg pair)
and, character 31.1 (solenomere flat). Despite being
closely related sister taxa, the distribution ranges of
the two genera include a 3000-km gap: Rondonaria is
limited to northern Brazil, whereas Odontopeltis is
limited to south-eastern Brazil.
The monophyly of the genus Odontopeltis is
strongly supported. The absence of a prefemoral
process, which is present in almost all members of the
family Chelodesmidae, is diagnostic. The presence of
the apical processes A and B in the gonopod is likely
to be unique within the family Chelodesmidae.
Gonopods of males in the genera Manfrediodesmus
(see Fig. 55) and Rondonaria (see Hoffman, 2006:
fig. 8) feature one or more apical processes. Whether
these are homologous to either process A or B in
Odontopeltis cannot be established at this time. A
wide-ranging comparative analysis of the male
gonopodal structure in Chelodesmidae is required to
define hypotheses of primary homology of these
processes. Within this data set the monophyly of
the genus Odontopeltis is well supported by four
nonhomoplastic synapomorphies: character 3.1
(suboval shape of the Tömösváry organ), character
15.1 (ozopore border not raised), character 25.1 (ectal
folds on telopodite shaft present), and character 39.1
(presence of a pair of macrobristles at the end of
prefemoral region and beginning of femoral region
of gonopods). Sister-group relationships within
Odontopeltis have some support: Clade A by character
14.2 (presence of three or more projections at the edge
of the seventh paranotum) and by character 38.2
(process B of the gonopod being dorsally directed);
Clade B by character 40.1 (wide spacing of the two
macrosetae on the telopodite shaft). Clade C is supported by character 33.1 (pointed median solenomere
projection) and character 35.2 (extension of the
solenomere base). Clade D, the sister-group relationship between Odontopeltis donabeja and Odontopeltis
xica is supported by character 38.2 (process B of
gonopod being posteriorly directed by reversal). Clade
E is supported only by homoplastic characters. This
particular data set confirms Hoffman’s (1980) choice
to remove Odontopeltis from tribe Telonychopodini.
Currently, Odontopeltis cannot be placed into the
existing tribal structure of the family Chelodesmidae.
A monophyletic tribe Macrocoxodesmini, as defined by
Golovatch & Hoffman (2004), is not supported in the
current data set: Eucampesmella appears to be the
sister to the Telonychopodini (supported by characters
21 and 43, Clade H, see above), whereas the sistergroup relationship of Macrocoxodesmus to Clade F is
well supported (characters 12, 16, and 22).
TAXONOMY
FAMILY CHELODESMIDAE COOK, 1895
SUBFAMILY CHELODESMINAE COOK, 1895
GENUS ODONTOPELTIS POCOCK, 1894
Rhacophorus Koch, 1847, 59: 137. Type species
Polydesmus conspersus Perty, 1833, by subsequent
designation of Pocock, 1909: 168. Pre-occupied by
Rhacophorus Kuhl & van Hasselt, 1822, Amphibia.
Neave, 4: 30; Jeekel, 1970: 284; Hoffman, 1980: 151.
Odontotropis Humbert & deSaussure, 1869: 152.
Type species Polydesmus (Odontotropis) clarazianus
Humbert & deSaussure, 1869, by monotypy. Preoccupied by Odontotropis Agassiz, 1846, Mollusca.
Attems, 1898: 408. Neave 3: 389, Neave 5: 180;
Jeekel, 1970: 276; Hoffman, 1980: 151.
Odontopeltis Pocock, 1894: 509. New name for
Rhacophorus. Silvestri, 1895a: 769, with type species
Polydesmus conspersus, by direct substitution, see
Rhacophorus. Silvestri, 1895b: 3; Pocock, 1909: 168;
Neave 3, 386; Jeekel, 1970: 275; Hoffman, 1980: 151;
Hoffman, 1981a: 55 (revalidation of the genus name).
Storthotropis Attems, 1938: 200. New name for
Odontotropis Humbert & deSaussure, 1869, with type
species Polydesmus clariazianus, by direct substitution, Odontotropis. Hoffman, 1980: 151 (synonymized
with Odontopeltis); Neave 5: 259; Schubart, 1949: 18;
Jeekel, 1970: 289.
Diagnosis: Males and females of Odontopeltis differ
from other chelodesmid genera by the oval-shaped
Tömösváry organ (character 3, Fig. 7A) and by
the flat ozopore rim (Fig. 14B). Males of the genus
differ from other Chelodesmidae by the following
unique characters: absence of prefemoral processes,
solenomere sickle-shaped (Fig. 4B) and laminar
(Fig. 4A, mesal view). Acropodite with two apical processes, process A and process B, in addition to the
solenomere; process B two times as long as process
A; two macrobristles demarcate distal end of the
prefemoral region (Fig. 4B); prefemoral region two to
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
755
three times as long as the femoral region; spermatic
groove straight in femoral region (mesal view),
turning ectally in the solenomere. Females differ from
those of all other genera in this data set, except
Rondonaria and Macrocoxodesmus marcusi (Macrocoxodesmus marcusi, vulva of female unknown),
by the short cyphopod, without apical elongation
(Fig. 53). This same feature is observed in females of
Rondonaria, which differ from Odontopeltis by the
posterior opening of the cyphopod in the former.
shaped, with a median projection (Fig. 43). Spermatic
opening with many internal spine-shaped projections
(Fig. 41).
Female
characters:
Cyphopods:
oval-shaped
cyphopod located right behind the second pair of legs
on the second body ring. Composed of two valves
united by a membrane (Figs 71–73), with a third
apically hinged sclerite, appressed against both lower
valves.
Description:
Body
length
between
80 mm
(Odontopeltis conspersus) and 107 mm (Odontopeltis
giganteus). Head: face with three rows of bristles
(Fig. 6). Antenna: distal antennomere with modified
bristles arranged in patches, bristles with a median
projection (Fig. 58); very small spines on the ectal
side. Body rings: integument with small tubercles on
the metazonite (Fig. 57), without bristles. Coloration:
animals preserved in alcohol range from black, red to
white. Living specimens black with yellow paranota
(Fig. 1). Collum: anterior border arched and posterior
border straight. Sternite of body rings 5 and 6 with
two or four ventral projections (Figs 17, 18), respectively. Ozopore arrangement: 5, 7, 9, 10, 12, 13, 15–19;
ozopores surrounded by peritremata. Paranota:
prominent and long, covering half of the legs. The
first four body rings, body rings 18 and 19 with simple
paranota without projections. Paranota of body
rings 5 to 17 with teeth-like projections (Figs 15, 16),
varying between the species. Body rings with
ozopores show a prominent projection owing to
the presence of peritrema. The species-specific
teeth-projection formula of the paranota follows
Schubart (1949), e.g.: 0-0-0-0-2-2-3-2-3-3-2-3-3-2-3-33-0-0 (Odontopeltis conspersus). Legs: pregonopodal
legs without modifications, but with higher concentration of bristles than on posterior legs. Telson: triangular, with five pairs of macrobristles on the dorsal
side and two pairs in the apical region.
Distribution: Known only from the south-eastern
region of Brazil, recorded from the states of Espírito
Santo, Minas Gerais and Rio de Janeiro.
Male characters: Gonopore: coxae of second pair of
legs ventrally elongated, sclerotized, the conical projection bearing an apical pore.
Gonopod opening on seventh body ring: trapezoidal,
with a depression on the posterior edge and a membranous support basis of the gonopod.
Gonopods: cylindrical, projecting anteriorly; coxae
with or without spiniform process. Spermatic groove
always on the mesal side of the gonopod. Prefemoral
region covered by bristles, with an angular basal
projection (BPPf). Prefemoral process absent.
Prefemoral region two or three times as long as
femoral region. Femoral region without bristles; with
three apical projections. Process B pointing dorsally;
process A varying in direction. Solenomere sickle-
Composition: Eight species, four previously described:
Odontopeltis conspersus (Perty, 1833), Odontopeltis
anchisteus Hoffman, 1981 (see Hoffman, 1981a),
Odontopeltis clarazianus (Humbert & deSaussure,
1869), Odontopeltis giganteus (Schubart, 1949), and
four new species: Odontopeltis tiradentes, Odontopeltis
xica, Odontopeltis aleijadinho, and Odontopeltis
donabeja.
SPECIES
INCERTAE SEDIS
Rhacophorus decoloratus C. L. Koch, 1847, type locality and deposition of type material unknown.
Note: according to Sierwald & Reft (2004), C. L.
Koch deposited specimens at five museums but we
were informed by their curators that type material
of Rhacophorus decoloratus was not located in any
of these: ZSM (Dr Roland Melzer), ZMB (Dr Jason
Dunlop), BMNH (Dr Janet Becalloni), ZIN (Dr Victor
A. Krivokhatsky), and ZMUZ (Dr Paul Ward).
Polydesmus gracilipes Humbert & deSaussure,
1870. Holotype female, from Rio de Janeiro, Brazil,
deposited in Naturhistorisches Museum Wien
(Vienna, Austria), examined. The specimen is not a
member of the genus but cannot be identified more
precisely.
Odontopeltis balzanii Silvestri, 1895, see Silvestri,
1895a: 769, fig 1. Holotype male, from Missiones
Mosetenes, Bolivia, should be in Museo Civico di
Storia Naturale, Genoa, Italy.
Note: the type was deposited in the Genoa Museum
and was loaned on 10 September 10 1902 to Filippo
Silvestri in Portici, Italy (Jeekel, 1965: 122). In 2005,
the Genoa Museum received the material back, but
the type specimen of Odontopeltis balzanii was not
returned (Dr Giuliano Doria, pers. comm.). Silvestri’s
(1895a) illustration shows a small prefemoral process,
and an acropodite with three apical projections.
The presence of the prefemoral process excludes this
species from the genus Odontopeltis.
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756
J. P. P. PENA-BARBOSA ET AL.
Odontopeltis borellii Silvestri, 1895, see Silvestri,
1895b: 3, fig 2. Holotype male, from Paraguay, deposited at MRSN, examined.
Note: in the same publication, Silvestri (1895b) also
described Strongylosoma borellii from Argentina on
page 5, accompanied by figure 7. In 1897, Silvestri
moved Strongylosoma borellii to Mestosoma Silvestri,
1897, erroneously quoting page 3 as the original
description, but clearly indicating the intention
of moving Strongylosoma borellii and not Odontopeltis borellii to Mestosoma. No other mention of
Odontopeltis borellii was found in the literature. The
type specimen does not belong to Odontopeltis as
evidenced by the presence of a prefemoral process on
the gonopod.
Odontopeltis proxima Silvestri, 1895, see Silvestri,
1895b: 5, fig. 6. Holotype male, from Argentina,
deposited at MRSN, examined.
Note: the gonopod of the specimen possesses
a well-developed ventrally projecting prefemoral
process, which covers the acropodite. Silvestri’s illustration of the gonopod is accurate. This feature
excludes the specimen from the genus Odontopeltis,
but identifies the specimen as a member of the family
Chelodesmidae. Silvestri’s drawing also indicates
torsion of the acropodite, which was not confirmed by
our examination of the type specimen.
ODONTOPELTIS CONSPERSUS (PERTY, 1833)
FIGURES 59–61, 86
Polydesmus conspersus Perty, 1833: 210, L, fig. 8.
Male holotype, from Minas Gerais, Brazil, deposited
at ZSM, lost (Dr Roland Melzer). Gervais, 1836: 378;
Gervais, 1837: 44; Brandt, 1839: 310; Lucas, 1840:
525; Gervais, 1847: 112.
Rhacophorus conspersus: – Koch, 1847: 137
(comb. nov.)
Rhacophorus conspersus: – Koch, 1863: 53, fig. 175.
Odontopeltis conspersus: – Pocock, 1894: 509 (comb.
nov.); Pocock, 1909: 168; Hoffman, 1981a: 56.
Neotype: Male from Betim (19°58′04′′S, 44°11′52′′W),
Minas Gerais, Brazil, without date or collector data,
here designated as neotype (IBSP 2852).
Note: the proposition of the neotype is based on the
original description by Perty (1833) and the description and figure by Koch (1863), who examined the
remains of Perty’s specimen. Perty (1833) gives
‘mountains of Minas Gerais’ as the type locality.
Additional material examined: BRAZIL. Minas
Gerais: Viçosa (20°45′14′′S, 42°52′55′′W), Campus da
Escola Superior de Agricultura, 1 male, 1940 (MZSP
906).
Figures 59–64. Gonopods of Odontopeltis, left gonopod
illustrated. Figures 59–61. Odontopeltis conspersus (Perty,
1833). Figure 59. Mesal view. PrB, process B. Figure 60.
Ventral view. Figure 61. Ectal view. Scale bar = 0.5 mm.
Figures 62–64. Odontopeltis anchisteus Hoffman, 1981.
Figure 62. Mesal view, arrow: fold on process A (character
37). Figure 63. Ventral view, arrow: median process
absent. Figure 64. Ectal view. Scale bar = 0.5 mm.
Diagnosis: Males of Odontopeltis conspersus differ
from all other species of Odontopeltis by the rounded
tip of process B (PrB), visible in mesal and ectal view
(Figs 59, 61). The shape of process A (PrA) is similar
to the one in Odontopeltis tiradentes, but is distinguishable by its oval shape (visible in mesal and
ventral view, Figs 74, 75). Odontopeltis tiradentes is
distinguishable from Odontopeltis conspersus by the
basal constriction of the solenomere in the former
(Fig. 75, arrow).
Redescription: Male (Neotype, IBSP 2852): Head red.
Body pale yellow to white, with many red spots in
dorsal view. Tips of paranota yellow or white. Legs
and telson yellow. Paranotal teeth formula: 0-0-0-0-22-3-2-3-3-2-3-3-2-3-3-3-0-0. Total length 90. Collum 5
long, 13 wide. Antennomere length: 2.2, 2.3, 2.3, 2.4,
?. Genital opening 2.1 long, 3.4 wide. Telson broken.
Gonopods: coxae with spiniform process, macrobristles vertically aligned, separated by less than
their length. Prefemoral region three times longer
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PHYLOGENY AND REVIEW OF ODONTOPELTIS
than femoral region. Process B subtriangular, pointing dorsally in ectal and mesal view (Figs 59, 61).
Process A oval (in ventral view, Fig. 60), without projections, half as long as solenomere. Solenomere with
a small median projection.
Female: Unknown.
Distribution: Minas Gerais, Brazil.
ODONTOPELTIS ANCHISTEUS HOFFMAN, 1981
FIGURES 62–64, 86
Odontopeltis anchisteus Hoffman, 1981: 58, figs 4–6,
10, 11. Male holotype, from Minas Gerais, Brazil, A.
Fry col., deposited at BMNH 1873.30, examined.
Additional material examined: BRAZIL: Minas
Gerais: Juiz de Fora, (21°45′50′′S, 43°21′00′′W), 1씹,
10.ix.2009 (IBSP 2874); Santa Fé de Minas,
(16°41′24′′S, 45°24′50′′W), 1씹, Koch col. (BMNH).
Diagnosis: Males differ from all other species of the
genus by the absence of a median process on the
solenomere (Fig. 63, arrow); the presence of a single
rounded projection on process A in ventral view
(Fig. 63) and an apical fold in mesal view (Fig. 62,
arrow).
Redescription: Male (holotype, BMNH 1873.30): Head
red. Body reddish. Paranota tip yellow. Legs brown.
Telson yellow. Paranotal teeth formula: 0-0-0-0-2-2-24-5-4-6-4-3-7-4-3-3-1-1. Total length: 90. Collum 4.5
long, 13.9 wide. Antennomere length: 2.2, 2.4, 2.5,
2.3, 2.1. Genital papilla 0.4 high, 0.68 wide. Genital
opening 2.0 long, 3.8 wide. Telson damaged. Gonopods:
coxae with spiniform process. Prefemoral region three
times longer than femoral region. Macrobristles short
and in an inclined line, distance between them equal to
their length. Process B projecting dorsally in mesal
and ectal view, distal half slightly curved (Fig. 64).
Process A with a fold (Figs 62, 64), visible in mesal and
ectal view. Solenomere without median projection
(Fig. 63, arrow) and a relatively long tapering tip.
Female: Unknown.
Distribution: Minas Gerais, Brazil.
ODONTOPELTIS CLARAZIANUS
(HUMBERT & DESAUSSURE, 1869)
FIGURES 65–67, 86
Polydesmus (Odontotropis) clarazianus Humbert &
deSaussure, 1869: 152. Female holotype from La
Plata, Argentina, deposited in MHNG, examined;
deSaussure & Humbert, 1872 (description of male).
Odontotropis clarazianus: – Attems, 1898: 408.
757
Storthotropis clarazianus: – Attems, 1938: 200
(comb. nov.), Schubart, 1954: 141.
Odontopeltis clarazianus: – Hoffman, 1981a: 57,
figs 1–3, 7, 9 (comb. nov.).
Note: this species was described based on a female
(Humbert & deSaussure, 1869) from La Plata,
Argentina. DeSaussure & Humbert (1872) mentioned a male, but neither designated this male as
a type nor presented a drawing. Brölemann (1900)
published two drawings of a male under the name
Odontopeltis clarazianus based on specimens collected by Emílio Goeldi in Serra dos Órgãos, Rio
de Janeiro (specimen lost, MNHN, J-.J. Geoffroy,
unpubl. data). Schubart (1949: 20, figs 1, 2) presented a drawing of a male identified by him as
Odontopeltis clarazianus from Matipó, Minas Gerais,
which he considered to be conspecific with the specimens illustrated by Brölemann (1900). However,
Hoffman (1981a) examined the female holotype of
Odontopeltis clarazianus from La Plata and concluded that the male discussed by deSaussure &
Humbert (1872) is conspecific with the female type
specimen, but not conspecific with the males
described by Brölemann (1900) and described and
figured by Schubart (1949). Based on similarities of
the body between the female type specimen and
male specimens, Hoffman selected a new male specimen as conspecific with the Odontopeltis clarazianus
female. Therefore, the male specimens described by
Brölemann (1900) and Schubart (1949) are neither
conspecific with Odontopeltis clarazianus nor with
any other described species of Odontopeltis. They
belong to a heretofore undescribed species, described
below as a new species: Odontopeltis tiradentes.
Additional material examined: BRAZIL: Rio de
Janeiro: Trajano de Morais, Povoado das Almas,
(22°03′46′′S, 42°03′57′′W), 1씹, 02.i.2010, P. D. P.
Pinheiro col. (IBSP). ARGENTINA: Departamento de
Buenos Aires: La Plata, 1씹 (BMNH) (designated by
Hoffman, 1981a: 57).
Diagnosis: Males of Odontopeltis clarazianus differ
from all other species of the genus by the presence
of three apical teeth-like projections on process A
(Fig. 66).
Redescription: Male (MHNG): Head and body reddish.
Paranotal tips yellow. Legs brown. Telson yellow.
Paranotal teeth formula: 0-0-0-0-3-5-4-7-5-6-6-3-3-44-3-3-1-1. Total length 98. Collum 5.0 long, 14.0 wide.
Antennae damaged. Genital papilla 0.9 high, 0.66
wide. Genital opening 1.0 long, 3.25 wide. Telson 2.5.
Gonopods: coxae with distinct spiniform process.
Macrobristles in an inclined line with the distance
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
758
J. P. P. PENA-BARBOSA ET AL.
Figures 71–73. Left cyphopod of Odontopeltis giganteus.
Figure 71. Posterior aspect. Figure 72. Anterior aspect.
Figure 73. Mesal view. Scale bar = 0.1 mm.
Odontopeltis giganteus: – Hoffman, 1981a: 60,
figs 8, 12.
Additional material examined: BRAZIL: 1씹 (without
label information) (MZSP 908); Espírito Santo: Santa
Teresa, (19°56′09′′S, 40°36′00′′W), 1씹 and 1씸, 19.i.
1964 (MZSP 564); Linhares, (19°23′27′′S, 40°04′19′′W),
2씹, 1981, B. Kuth coll. (MNRJ 11950).
Figures 65–70. Gonopods of Odontopeltis, left gonopod
illustrated. Figures 65–67. Odontopeltis clarazianus
(Humbert & deSaussure, 1863). Figure 65. Mesal view.
Figure 66. Ventral view. Figure 67. Ectal view. Scale
bar = 0.5 mm. Figures 68–70. Odontopeltis giganteus
(Schubart, 1949). Figure 68. Mesal view, arrow: posterior
hump of solenomere. Figure 69. Ventral view, arrow:
median process of solenomere. Figure 70. Ectal view. Scale
bar = 0.5 mm.
between them equal to their length. Prefemoral
region 2.5 × longer than femoral region. Process B is
slightly bent in clockwise direction when observed in
ventral and mesal view (Figs 65, 66). Process A
subrectangular, with three apical teeth-like processes
(Fig. 66). Solenomere with a small median projection.
Female (holotype, MHNG): Body colour as in male.
Paranotal teeth formula: 0-0-0-0-3-3-2-3-2-3-3-2-2-34-2-2-1-1. Total length 96. Collum 4.7 long, 13.1
wide. Antennae damaged. Telson 2.1. Genitalia as in
Odontopeltis giganteus (Figs 71–73).
Distribution: Rio de Janeiro, Brazil, and La Plata,
Argentina.
ODONTOPELTIS GIGANTEUS (SCHUBART, 1949)
FIGURES 68–73, 86
Storthotropis giganteus Schubart, 1949: 22, figs 3, 4.
Male holotype, from Espírito Santo, Brazil, ii.1911, E.
Garbe coll., deposited in MZSP 100, examined.
Diagnosis: Males of Odontopeltis giganteus differ from
all other species of the genus by the curved tip of
process B (Fig. 68). Process B large, not obscured by
solenomere in ventral view (Fig. 69). Coxae without
spiniform process (Fig. 68). Femoral region as wide
as prefemoral region, unlike in all other species of
Odontopeltis.
Redescription: Male (Holotype, MZSP 100): Head red.
Body reddish. Paranotal tips and telson yellow. Legs
reddish. Paranotal teeth formula: 0-0-0-0-2-1-2-2-3-33-2-3-3-2-2-2-1-0-0. Total length 107. Collum 5.1 long,
15.6 wide. Antennae missing. Genital opening 2.5 long,
4.08 wide. Telson 1.4. Gonopods: macrobristles in an
inclined line, close together, distance between them
less than half length of a bristle. Prefemoral region
2.5 × longer than femoral region. Process A triangular
and long, almost the size of the solenomere, without
projections. Solenomere with a median process (arrow,
visible in ventral view, Fig. 69) and with a hump
(arrow, visible in mesal view, Fig. 68).
Female: Body as in male. Paranotal teeth formula:
0-0-0-0-2-2-3-2-3-2-2-3-3-2-2-2-1-1-1. Total length 89.
Collum 5.0 long, 14 wide. Telson 2.0. Cyphopod short
and oval-shaped, covered by bristles. Ectal valve
slightly bigger than mesal valve (Figs 53, 71). Mesal
valve subrectangular (mesal view, Fig. 73). Ectal
valve lozenge-shaped (Fig. 72).
ODONTOPELTIS TIRADENTES
FIGURES 74–76, 87
SP. NOV.
Storthotropis clarazianus: Schubart, 1949: 20, figs 1,
2 (male description, from MZSP, misidentification);
Brölemann, 1900: 65 (male = Odontopeltis tiradentes).
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
PHYLOGENY AND REVIEW OF ODONTOPELTIS
759
Additional material examined: BRAZIL: Minas
Gerais: Matipó, Fazenda Floresta, (20°16′S, 42°21′W),
only male gonopods from Schubart, viii.1919, P. da
Fonseca coll. (MZSP 909).
Etymology: The species epithet is a noun in apposition, after the nickname Tiradentes of Joaquim José
da Silva Xavier, a Brazilian national hero known for
leading the Inconfidência Mineira in 1789.
Diagnosis: Males of Odontopeltis tiradentes are
similar to Odontopeltis giganteus in the slightly
curved tip of process B in mesal view (Fig. 74).
Odontopeltis tiradentes differ from Odontopeltis
giganteus by the wider shape of process A (Fig. 75,
ventral view), and by the absence of a hump on the
solenomere (in mesal view, Fig. 74). Odontopeltis
tiradentes differs from all other species of the genus
by a constriction in the basis of the solenomere
(Fig. 75, ventral view, arrow).
Figures 74–79. Gonopods of Odontopeltis, left gonopod
illustrated. Figures 74–76. Odontopeltis tiradentes sp.
nov. Figure 74. Mesal view. Figure 75. Ventral view,
arrow: constriction in base of solenomere. Figure 76. Ectal
view. Scale bar = 0.5 mm. Figures 77–79. Odontopeltis
donabeja sp. nov. Figure 77. Mesal view. Figure 78.
Ventral view, arrow: depression on process A. Figure 79.
Ectal view. Scale bar = 0.5 mm.
Type material: Male holotype from Viçosa, (20°45′
14′′S, 42°52′55′′W), Minas Gerais, Brazil, 1947, deposited in MZSP, catalog number 909; labelled as
‘O. clarazianus?’, most likely one of the specimens
Schubart examined.
Note: this species is based on a male identified
as Odontopeltis clarazianus by Schubart (1949),
who based his identification on gonopod drawings
by Brölemann (1900). Hoffman (1981a) noted that
Brölemann’s and Schubart’s specimens are not
conspecific with the female type specimen of
Odontopeltis clarazianus. The original specimen
described by Brölemann is lost, and so is the body of
Schubart’s specimen. However, the gonopods of
Schubart’s specimen, mounted on microscope slides,
are available (MZSP 909), and were compared to
another specimen in MZSP labelled Odontopeltis
clarazianus, now holotype of Odontopeltis tiradentes.
The diagnosis and description of Odontopeltis
tiradentes are based on these two specimens.
Description: Male (holotype): Head and body red. Tip
of paranota and telson yellow. Legs orange. Paranotal
teeth formula: 0-0-0-0-2-4-3-4-3-3-5-3-3-4-3-3-3-1-1.
Total length 90. Collum 5 long, 14 wide. Antennae:
2.4, 2.0, 2.0, 2.1, 2.1. Genital opening 3.9 wide, 2.4
long. Telson 2.0. Gonopod coxa with a spiniform
process. Prefemoral region 3 × longer than femoral
region. Macrobristles on an inclined line, close
together, with the distance between them less than
half the length of the bristle itself. Process B with
the tip slightly curved ectally. Process A elongated,
without projections. Solenomere with a basal constriction and a well-developed median projection.
Female: Unknown.
Distribution: Minas Gerais, Brazil.
ODONTOPELTIS DONABEJA SP.
FIGURES 77–79, 87
NOV.
Type material: Male holotype from Estação Ecológica
de Peti, CEMIG, São Gonçalo do Rio Abaixo
(19°49′33′′S, 43°21′43′′W), Minas Gerais, Brazil,
03−05.x.2008, I. S. Oliveira coll., deposited in IBSP
1668.
Paratype: Female from the same locality, 05−08.i.
2010, J. P. P. Pena-Barbosa et al. coll. (IBSP 3618).
Additional material examined: BRAZIL: Minas
Gerais: São Gonçalo do Rio Abaixo, (19°49′33′′S,
43°21′43′′W), Estação Ambiental de Peti, CEMIG, 7씹,
4씸, 05−08.i.2010, J. P. P. Pena Barbosa et al. coll.
[IBSP 3617 (1씸), 3620–3621 (2씸), 3622–3625 (4씹),
3626 (1씸), 3627 (1씹), 3629 (1씹), 3693 (1씹)].
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
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J. P. P. PENA-BARBOSA ET AL.
Etymology: The species epithet is a noun in apposition, a patronym in honour of Ana Jacinta de São
José, known as Dona Beja, a woman from Minas
Gerais known for her beauty and curious life history.
Diagnosis: Males of Odontopeltis donabeja differ from
all other species of the genus by the large and distinct
median process of the solenomere, surpassing the tip
of the solenomere (Figs 78, 79). Males of Odontopeltis
xica are similar with regard to the posteriorly curved
tip of process B; however, the curvature is much more
pronounced in Odontopeltis xica. In Odontopeltis
donabeja process B is subtriangular in shape,
whereas it is rounded in Odontopeltis xica.
Description: Male (holotype, IBSP 1668): Head and
body black. Tip of paranota and telson yellow. Legs
brown. Paranotal teeth formula: 0-0-0-0-2-2-3-2-3-32-4-3-2-3-3-3-1-1. Total length 87. Collum 5 long, 13
wide. Antenna: 2.3, 2.2, 1.9, 2.0, 2.3. Genital opening
2.0 long, 3.5 wide. Telson 1.9. Gonopod coxa with a
spiniform process. Prefemoral region 2.5 × longer
than femoral region. Macrobristles on an inclined line
with the distance between them less than half the
length of the bristle itself. Process A with a depression
(Fig. 78, arrow). Process B subtriangular and slightly
curved to posterior.
Female (paratype, IBSP 3618): Coloration as in
male. Paranotal teeth formula as in male. Total
length 80. Collum 4 long, 12 wide. Antenna: 2.1, 1.7,
1.9, 2.1, 2.0.
Figures 80–85. Gonopods of Odontopeltis, left gonopod
illustrated. Figures 80–82. Odontopeltis aleijadinho
sp. nov. Figure 80. Mesal view. Figure 81. Ventral view.
Figure 82. Ectal view. Scale bar = 0.5 mm. Figures 83–85.
Odontopeltis xica sp. nov. Figure 83. Mesal view.
Figure 84. Ventral view. Figure 85. Ectal view. Scale
bar = 0.5 mm.
Distribution: Minas Gerais, Brazil.
ODONTOPELTIS ALEIJADINHO
FIGURES 80–82, 87
SP. NOV.
Type material: Male holotype from Parque Estadual
do Rio Doce, Timóteo (19°34′58′′S, 38°38′W), Minas
Gerais, Brazil, 17−21.xi.2008, I. S. Oliveira coll.,
deposited in IBSP, catalog number 1542.
Paratype: One male (IBSP 2853) and one female
(IBSP 3630) from the same locality.
Etymology: The species epithet is a noun in apposition,
a patronym in honour of Antônio Francisco Lisboa,
nickname Aleijadinho, one of the most important
artists of the Baroque, and who came from Minas
Gerais.
Diagnosis: Males of Odontopeltis aleijadinho differ
from other species of the genus by the rounded tip
of process B. In both Odontopeltis aleijadinho and
Odontopeltis anchisteus process A possesses a fold: in
Odontopeltis anchisteus this fold is positioned api-
cally, whereas in Odontopeltis aleijadinho the fold is
at the base (Figs 80, 82). Odontopeltis tiradentes and
Odontopeltis aleijadinho both possess a subtriangular
process A with small apical undulations.
Description: Male (holotype, IBSP 1542): Head and
body black. Paranota tip and telson yellow. Legs black.
Paranotal teeth formula: 0-0-0-0-2-2-3-3-3-3-5-3-3-3-43-3-1-1. Total length 86. Collum 4 long, 13 wide.
Antenna: 2.3, 2.0, 2.0, 2.0, 2.5. Genital opening 1.9
long, 3.3 wide. Telson 1.8. Coxae of gonopods with a
spiniform process. Prefemoral region 3 × longer than
femoral region. Macrobristles on an inclined line with
the distance between them less than half the length of
the bristle itself. Process A without apical process, only
with small ripples (ventral view, Fig. 81). Median
projection of the solenomere well developed, but not
surpassing tip of solenomere (Fig. 81).
Female (paratype, IBSP 3630): Coloration as in
male. Paranotal teeth formula: 0-0-0-0-2-2-3-2-3-3-23-3-2-3-3-2-1-1. Total length 87. Collum 5 long, 12
wide. Antenna: 2.2, 2.0, 2.0, 1.9, 1.9. Telson 1.8.
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
PHYLOGENY AND REVIEW OF ODONTOPELTIS
761
Figures 86, 87. Distribution map of Odontopeltis species. Figure 86. States: Rio Grande do Sul to Espírito Santo:
Odontopeltis anchisteus, Odontopeltis clarazianus, Odontopeltis conspersus, and Odontopeltis giganteus (Espirito Santo).
Figure 87. State: Minas Gerais: Odontopeltis tiradentes sp. nov., Odontopeltis donabeja sp. nov., Odontopeltis
aleijadinho sp. nov., and Odontopeltis xica sp. nov.
Distribution: Know only from the type locality.
ODONTOPELTIS XICA SP.
FIGURES 83–85, 87
NOV.
Type material: Male holotype from Parque Nacional
da Serra do Cipó, Santana do Riacho, (19°57′52′′S,
43°24′54′′W), Minas Gerais, Brazil, deposited in IBSP
1562.
Etymology: The species epithet, a noun in apposition,
is a patronym in honour of Francisca da Silva de
Oliveira, known as Xica da Silva, a famous slave
woman who became extremely rich in Minas Gerais.
Diagnosis: Males of Odontopeltis xica differ from all
other species of the genus by the small size of process
A (Fig. 83). The tip of process B is curved posteriorly
(Figs 83, 85).
Description: Male (Holotype, IBSP 1562): Head, body
and legs blackish. Paranota tip and telson yellow.
Paranotal teeth formula: 0-0-0-0-2-2-3-2-3-3-2-4-3-23-3-1-0-0. Total length 86. Collum 4 long, 13 wide.
Antenna: 2.3, 2.0, 2.0, 2.0, 2.5. Genital opening 1.9
long, 3.3 wide. Telson 1.8. Prefemoral region 3 ×
longer than femoral region. Macrobristles on the
same plane and with distance between them less than
half the length of the bristle itself. Coxae of gonopods
with a small spiniform process. Process A small and
suboval, without apical process. Median process of the
solenomere well-developed, projecting apically, not
surpassing the tip of the solenomere.
Female: Unknown.
Distribution: Known only from the type locality.
ACKNOWLEDGEMENTS
The authors are grateful to the institutions and their
curators for the loan of specimens for this work, and
to A. B. Kury (MNRJ), A. J. Santos (UFMG), and R.
Pinto da Rocha (MZSP) for their hospitality during
visits. The first author is grateful to B. A. S. Medeiros
(Harvard University) for his comments and help
with the implied weighting techniques. He is also
grateful to I. L. F. Magalhães (UFMG) for discussion
of the study, help with specimen collection, and for
generating the maps. Thanks are due to the staff of
the Centro de Microscopia Eletrônica d IB-USP for
assisting with the scanning electron microscope and
generating the SEM images. We greatly appreciate
the valuable comments on previous drafts of the
manuscript by A. Chagas-Jr (UFMT) and A. J. Santos.
© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 169, 737–764
762
J. P. P. PENA-BARBOSA ET AL.
Furthermore, we are grateful for the careful reviews
of the manuscript by Dr Henrik Enghoff and an
anonymous reviewer, initiated by the editor Dr Peter
Hayward. This study was financially supported by a
CNPq scholarship to J. P. P. Pena Barbosa (grant
number 130380/2009-1), by a grant from FAPESP
(process number 2009/10343-7) to A. D. Brescovit, and
by visiting scholarship grants of the Field Museum to
J. P. P. Pena Barbosa.
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