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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4061 (3): 253
–
260
http://www.mapress.com/j/zt/
Article
253
http://doi.org/10.11646/zootaxa.4061.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:F438B8B7-F411-4998-9D62-B0DAEADE3B3B
Morphological tricks and blessed genitalia: rectifying the family placement of
Fijicolana tuberculata (Opiliones: Laniatores: Zalmoxidae)
ABEL PÉREZ-GONZÁLEZ
1, 3
, PRASHANT P. SHARMA
2
& DANIEL N. PROUD
1
1
División Aracnología, Museo Argentino de Ciencias Naturales, CONICET, Av. Ángel Gallardo 470, C1405DJR Buenos Aires,
Argentina
2
Department of Zoology, University of Wisconsin-Madison, Madison, WI 53706,
USA
3
Corresponding author. E-mail: abelaracno@gmail.com
Abstract
The type specimens of Fijicolana tuberculata Roewer, 1963 were re-examined and the male genital morphology is illus-
trated and described for the first time. Despite the presence of several morphological features that are typical of Samoidae,
such as the presence of scopulae on legs III and IV, genital morphology unambiguously indicates that this species belongs
to the Zalmoxidae rather than to the Samoidae. Fijicolana Roewer, 1963 is newly synonymized with Zalmoxis Sørensen,
1886. However, the newly implied combination is preoccupied by Z. tuberculatus Goodnight & Goodnight, 1948 thus the
replacement name Zalmoxis roeweri nom. nov. is proposed to avoid secondary homonymy. The definition of Z. roeweri
nom. nov. is amended, and the morphology of this species is compared with other representatives of Zalmoxidae and Sam-
oidae. We conclude that the presence of scopulae alone is not a sufficiently diagnostic characteristic for Samoidae and,
therefore, correctly placing taxa into families within Samooidea + Zalmoxoidea requires additional morphological evi-
dence (e.g. genital morphology). In light of this result, we point out that the "scopulated" Australasian samoids Badessania
metatarsalis Roewer, 1949, Sawaiellus berlandi Roewer, 1949 and Parasamoa gressitti Goodnight & Goodnight, 1957
require re-examination in order to detect potential errors in their family placement.
Key words: Samoidae, taxonomy, scopula, penis morphology, Fiji, Melanesia
Introduction
In the highly criticized Roewerian system (e.g. Giribet & Kury 2007) the presence of a dense tarsal scopula on legs
III and IV is a crucial character to recognize subfamilies (many of them since elevated to families) in Roewer’s
Phalangodidae concept. In this typological approach the presence of scopulae was used to separate Samoinae from
Phalangoninae/Tricommatinae, Ibaloniinae from Podoctinae/Erecananinae and Acrobuniinae from Epedaninae
(Roewer 1912). Adhering to this schema until his last published work, Roewer (1963) created one more monotypic
genus by describing a new species from the Fiji Islands: Fijicolana tuberculata Roewer, 1963. This species
exhibits several features typical of Samoidae (see Pérez-González & Kury 2007), such as an incrassate metatarsus
III of the male and a remarkably dense tarsal scopula on legs III and IV in both sexes, key characters that led
Roewer to place this taxon in the Samoinae. A re-examination of the type series of F. tuberculata, and in particular
the study of male genital morphology, allowed us to clarify the family placement of this species, provide additional
taxonomic characters, and propose nomenclatural changes.
Material and methods
Type specimens of Fijicolana tuberculata deposited in the SMF (Senckenberg Naturmuseum und
Forschungsinstitut, Frankfurt, Germany) were examined. Other specimens used for morphological comparison are
deposited in the MCZ (Museum of Comparative Zoology, Harvard University, Cambridge, USA) and in the INBIO
(Instituto Nacional de Biodiversidad de Costa Rica, Santo Domingo, Heredia, Costa Rica).
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Photographs of ethanol preserved specimens were taken with a Leica DFC 290 digital camera attached to a
Leica M165C stereomicroscope, and different focal planes were combined using Helicon Focus Pro
(www.heliconsoft.com). Male genitalia preparation followed Acosta et al. (2007), with temporary mounts
embedded in glycerol. Male leg III was temporarily mounted and cleared in clove oil. Penis and leg III of the male
were drawn using a camera lucida attached to an Olympus BH-2 compound microscope. Genitalia drawings were
digitized using Corel Draw X7. Figures were edited using Photoshop CS5 or Corel Draw X7. For SEM preparation
the body parts were dehydrated in a series of increasing concentrations of ethanol (85%, 90%, 95%, 100%), and
air-dried before being fixed on a stub and sputter-coated. Micrographs for Badessa ampycoides were obtained
using the protocol described in Gainett et al. (2014).
Taxonomy
Zalmoxidae Sørensen, 1886
Zalmoxis Sørensen, 1886
Zalmoxis Sørensen, 1886: 64 [type species: Zalmoxis robusta Sørensen, 1886, by subsequent designation in Roewer 1949].
Fijicolana Roewer, 1963: 223 [type species: Fijicolana tuberculata Roewer, 1963, by original designation] syn. nov.
See the remainder of the extensive synonymy in Sharma et al. (2011: 42–43).
Justification of synonymy. Our decision follows the same rationale proposed by Sharma et al. (2011), based on
evidence from a molecular phylogeny which indicates that the Indo-Pacific Zalmoxidae constitute a monophyletic
group (Sharma & Giribet 2012), thus disfavoring monotypic genera that would render Zalmoxis para- or
polyphyletic. The habitus of F. tuberculata is zalmoxid-like, with a pyriform dorsal scutum, and remarkably similar
to that of the New Caledonian species Zalmoxis neocaledonicus Roewer, 1912 and Zalmoxis mendax Sharma,
2012. Additionally, the well-marked bulla, male genital morphology, and enlarged metatarsus III are very similar to
other Zalmoxis species (see below). The only two morphological features discordant with the most recent diagnosis
of Zalmoxis (see Sharma et al. 2011) are the presence of five tarsomeres on leg IV (rather than six in typical
Zalmoxis species) and the presence of tarsal scopulae on legs III and IV. Sharma et al. (2011) stated that the tarsal
formula for Zalmoxis is 3: 5–9: 5: 6, however a re-examination of all tarsal formulae described for the species
currently placed in Zalmoxis shows greater variation in all legs: 3–4: 4–9: 4–5: 5–6 (Table 1). The presence of five
tarsomeres on leg IV, supposedly diagnostic for Fijicolana, is currently shared by 11 other Zalmoxis species (Table
1). Therefore the presence of a scopula remains as the only diagnostic feature of Fijicolana that would separate it
from other species of Zalmoxis. Given that molecular evidence supports the monophyly of Zalmoxis in the Pacific
Islands, and that scopulae on tarsi III and IV are recurrently observed in unrelated lineages (e.g. Ibaloniinae; Kury
2007, Sharma & Giribet 2011), the scopula alone is insufficient to support F. tuberculata as belonging to a genus of
its own and instead is regarded as a putative autapomorphy of this species.
Zalmoxis roeweri nom. nov.
(Figs 1, 2A–C, 3A–E)
Fijicolana tuberculata Roewer, 1963: 223, pl. 18, figs 1–4.
Zalmoxis tuberculatus (Roewer, 1963) comb. nov., by this act becoming a junior secondary homonym of Zalmoxis tuberculatus
Goodnight & Goodnight, 1948).
Placement. Originally in the Phalangodidae: Samoinae. Here transferred to the Zalmoxidae.
Type material. Male holotype, one male and one female paratypes from MELANESIA, Fiji; SMF 9911155-
RII/11155-3; examined. Remark: The paratypes are one male and one female instead of two females as stated by
Roewer (1963) in the original description. The male paratype is poorly preserved and has presumably dried up in
the past.
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TABLE 1. Species of Zalmoxis with tarsomere counts that differ from the most common tarsal formula 3: 5–9: 5: 6
(deviations are shown in bold).
FIGURE 1. Zalmoxis roeweri nom. nov. (previously Fijicolana tuberculata Roewer, 1963), male holotype (A, C–F), female
paratype (B). (A) left metatarsus and tarsus III, lateral view. (B) left metatarsus III, lateral view (C–D) habitus, dorsal view. (E)
left femur and patella IV, lateral view. (F) habitus, lateral view. Scale bars 1 mm.
Species I II III IV
Zalmoxis bendis Sharma et al., 2012 3 6 4–5 5
Zalmoxis brevipes (Roewer, 1949) 3 5 5 5
Zalmoxis crassitarsis Suzuki, 1982 3 6 5 5
Zalmoxis cuspanalis (Roewer, 1927) 3 6 5 5
Zalmoxis dammermani (Roewer, 1927) 3 5 5 5
Zalmoxis derzelas Sharma et al., 2012 3 6 5 5
Zalmoxis gebeleizis Sharma et al., 2012 3 6 5 5
Zalmoxis kaktinsae Sharma, 2012 3 455
Zalmoxis lavacaverna Hunt, 1993 3 5 5 5
Zalmoxis mendax Sharma, 2012 475 6
Zalmoxis mitobatipes (Roewer, 1926) 3 6 5 5
Zalmoxis roeweri nom. nov. 37 5 5
Zalmoxis similis Suzuki, 1982 3 6 5 5
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FIGURE 2. (A–C) Zalmoxis roeweri nom. nov. (previously Fijicolana tuberculata Roewer, 1963), male paratype. (D–E)
Badessa ampycoides Sørensen, 1886 (Samoidae) from Fiji, male; pictures courtesy of Guilherme Gainett and Gonzalo Giribet .
(A) left tarsus III, lateral view. (B) distal portion of left metatarsus III, lateral view. (C) patella to tarsus of left leg III, lateral
view. (D) distitarsus III, dorsal view, setae removed. (E) metatarsus III, ventral view. Scale bars: A, B: 0.2 mm; C: 0.4 mm; D:
10 µm; E: 100 µm. As: astragalus; Ca: calcaneus; Di: distitarsus; DL: distitarsal lobes; Met: metatarsus; Pa: patella; Sc:
scopula; Ta: tarsus; Ti: tibia.
Etymology. The new name is a patronym in honor of the German arachnologist Carl Friedrich Roewer who
first studied and described this species.
Diagnosis. Distinguished from other Zalmoxis species by the presence of a conspicuous, dense scopula on the
terminal tarsomere of legs III and IV in both sexes; five tarsomeres on leg IV; metatarsus III of males incrassate;
anal operculum unarmed. Additionally, it is distinguished from Zalmoxis derzelas Sharma et al., 2012 by larger
body size and differences in armature of femur IV and male genital morphology, particularly the very wide rutrum.
Genital morphology. Penis (Figs 3A–E): Pars distalis well-defined, wider than pars basalis. Pergula and
rutrum clearly distinguishable. Rutrum very wide, apically shaped like an arrowhead, only slightly wider than base
(Fig. 3C). Setae above pergula arranged in three medial ventral pairs and two lateral groups of three. Dorsal-most
seta in each lateral group larger than others. One pair of small setae widely separated from each other, situated
ventrolaterally below pergula. Capsula externa modified into a massive stragulum, basally fused, with two small
dorso-subapical projections. Capsula interna entirely concealed by stragulum (in unexpanded penis), with a long,
thin stylus and two rigid, laminar, curved conductors.
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FIGURE 3. (A–E) Zalmoxis roeweri nom. nov. (previously Fijicolana tuberculata Roewer, 1963), penis of male paratype. (F–
G) Ethobunus zalmoxiformis (Roewer, 1949), SEM micrographs of penis parts. (A) distal portion of penis, dorsal view. (B)
same, lateral view. (C) same, ventral view. (D) same, dorsal view, showing internal morphology. (E) same, lateral view,
showing internal morphology. (F) capsula externa, lateral view. (G) capsula interna, laterodorsal-apical view. Scale bars: A–E
(to same scale): 100 µm; F, G: 100 µm. CE: capsula externa; CI: capsula interna; Co: conductor; DE: ductus ejaculatorius; PB:
pars basalis; PD: pars distalis; Pe: pergula; Ru: rutrum; S: stylus; St: stragulum.
Discussion
The family Samoidae was erected by Sørensen (1886) to include four species in three genera from the Pacific
Islands: Badessa ampycoides Sørensen, 1886, Samoa obscura Sørensen, 1886, Samoa variabilis Sørensen, 1886,
and Feretrius quadrioculatus (L. Koch, 1865). Sørensen (1886) took great care in defining this group on the basis
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of a broader morphological characterization, but he also used the presence of a tarsal scopula on legs III and IV of
males as a distinctive characteristic for his new family. A careful review of the literature and an examination of
some type specimens (unpublished data) indicate that these four species are closely related, and given that Samoa
Sørensen, 1886 is the type genus of the family, these taxa represent the typical Samoidae as defined by Pérez-
González & Kury (2007). It should be noted, however, that the highest samoid diversity is found in the Neotropics,
and the close ties between Neotropical and Australasian samoids is supported by morphological (unpublished data)
and molecular phylogenetic evidence (Sharma & Giribet 2011). The disjunct amphi-Pacific distribution of samoids
is similar to that of zalmoxids, the latter of which probably arrived in Australasia by transoceanic dispersal out of
the Neotropics (Sharma & Giribet 2012).
Decades after Sørensen (1886) there was a resurgence of Laniatores studies, and authors revisited the
systematics of the Pacific Island samoids, adding the species: Badessania metatarsalis Roewer, 1949, Sawaiellus
berlandi Roewer, 1949, Waigeucola palpalis Roewer, 1949, F. tuberculata, Parasamoa gressitti Goodnight &
Goodnight, 1957 and Zalmoxista australis (Sørensen, 1886) (see Roewer 1949 and 1963, Goodnight & Goodnight
1957, Pérez-González & Kury 2007). Although Sørensen, Roewer, C. J. Goodnight and M. L. Goodnight had not
recognized genital morphology as an important source of systematic characters, Sørensen was much more
meticulous about detailed morphological characteristics that defined the family. The other authors were overly
influenced by two features that exhibit extraordinary evolutionary convergence: the tarsal scopula and the
incrassate metatarsus III exhibited by males of some species.
The male genitalia unambiguously support the inclusion of Z. roeweri nom. nov. in the Zalmoxidae. The penis
morphology corresponds to the groundplan described for the family (see Kury & Pérez-González 2007) and it is
very similar to other Southeast Asian and Australasian species (see Sharma 2012, Sharma et al. 2012). The capsula
interna morphology is widely unstudied for zalmoxids, despite its potential for providing valuable diagnostic
characters. We did not expand the penis of Z. roeweri nom. nov. due to the scarcity of available specimens (only
two males exist in the type series), but we were able to observe the presence of two rigid and curved laminar
conductors and of a long, thin stylus (Figs 3D–E). A similar morphology of the capsula interna could be observed
in Ethobunus zalmoxiformis (Roewer, 1949) from Costa Rica (Volcán Poás National Park, Alajuela Province,
25.x.2004, A. Pérez-González, C. Víquez, J. Mata, R. Gutierrez, INBIO). Nevertheless, the capsula externa is
greatly different in both species (Figs 3F–G). The presence of two laminar conductors and of a long and thin stylus
is probably shared across the Zalmoxidae, but further studies are needed. Unfortunately, the genital morphology of
other Fijian Zalmoxidae [Zalmoxis insularis (Roewer, 1949), Zalmoxis pygmaeus Sørensen, 1886 and Zalmoxis
robustus Sørensen, 1886] remains unknown, thus preventing comparisons of Z. roeweri nom. nov. with the
Zalmoxis species recorded from the islands of Fiji. External characters such as the well-marked bulla, pyriform
body shape, and sexually dimorphic leg IV also support the synonymy of Fijicolana with Zalmoxis.
Two of the most remarkable (and tricky) morphological features of Z. roeweri nom. nov. are the incrassate
metatarsus III (in males; Fig. 1A cf. Fig. 1B) and the scopulae on legs III and IV (in both sexes; Fig. 2A, C)
because they are most commonly exhibited by members of Samooidea. After careful examination we detected that
the metatarsus III of Z. roeweri nom. nov. males is only superficially similar to that of samooid harvestmen (Fig.
2E). The samooidean metatarsus III possesses a long intrusion of the astragalus by the calcaneus on the ventral
surface, thus forming a very peculiar structure with glandular pores and modified setae (presumably with a
secretory function, see Pérez-González & Kury 2007, Willemart et al. 2010; contra Šilhavý 1979 who advocates
for a stridulatory function). The "samooidean type" of metatarsus III is present in B. ampycoides, a samoid from
Fiji (Fig. 2E), but it is absent in Z. roeweri nom. nov. where the swelling is restricted to the astralagus and where
there is no ventral calcaneus intrusion (Fig. 2B). This different kind of incrassate metatarsus III also occurs in other
Zalmoxis species such as Z. derzelas, Zalmoxis gebeleizis Sharma et al., 2012, and Zalmoxis zibelthiurdos Sharma
et al., 2012 (Sharma et al. 2012: figs 3e, 7e, 13e).
The scopulae on distitarsi of legs III and IV of Z. roeweri nom. nov. are dense and very well developed (Fig.
2A), remarkably similar to scopulae observed in samooidean taxa. We were not able to examine the legs under the
scanning electron microscope, but under the compound microscope it appears that the scopula is composed of the
same kind of apically enlarged setae that are present in several species of Samoidae (see Šilhavý 1979: fig. 3 and
unpublished data) as well as in some other distant families (see Pinto-da-Rocha 1997: fig. 587; Rambla 1990). One
major difference is present in the form of the apical surface of the distalmost tarsomere. In Samoidae the end of the
distalmost tarsomere is ventrolaterally projecting forward, thus forming two lobes on which ventral scopular hairs
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are situated basally. These scopulate tarsal lobes extend beyond the base of the claw, forming a deep cavity (see
Šilhavý 1979: fig. 2). This kind of distitarsus morphology can be seen in the Fijian samoid B. ampycoides (Fig.
2D). In Z. roeweri nom. nov., however, the distalmost tarsomere lacks such lobes (Fig. 2A).
Given that the samoid-like characteristics exhibited by Z. roeweri nom. nov. are only superficially similar to
those present in samooidean harvestmen, and that they probably evolved convergently in different lineages, we
conclude that the presence of a scopula is not diagnostic for Samoidae and that incrassate leg segments need to be
studied and compared in greater detail than before. Additionally, when placing taxa into families belonging to the
clade Samooidea + Zalmoxoidea, an examination of their genital morphology is crucial. As the presence of a
scopula has been a key characteristic for family assignment in the past, a re-evaluation of poorly described samoids
(in particular Australasian species) is needed to eliminate taxonomic misplacements.
We have demonstrated that the placement of species in the Samoidae on the basis of a scopula and of a slightly
incrassate metatarsus III in males is problematic. The transfer of Fijicolana from the Samoidae to the Zalmoxidae
points out only a single instance of taxonomic misplacement by early authors stemming from the highly convergent
morphology of these two closely related families. We suspect that other Australasian “samoids” may also belong to
the Zalmoxidae or possibly to other families. Although we have not yet examined all type specimens of the other
Australasian samoids, our study casts doubt on the placement of the "scopulated" species Badessania metatarsalis,
Sawaiellus berlandi and Parasamoa gressitti. The strongly developed basichelicerite with a well-marked bulla
illustrated in the original descriptions of these species raises doubts about their current placement in the Samoidae
and suggests inclusion in the Zamoxidae. It will be necessary to carry out a detailed study of the external and
genital morphology of these species. The present work, together with the recent transfer of Waigeucola palpalis to
Podoctidae (Pérez-González 2011), improves our understanding of the systematics of Samoidae, Zalmoxidae and
Podoctidae. Interestingly, this work has revealed more about Samoidae than Zalmoxidae, the latter having already
been thoroughly examined across the Pacific Islands (Sharma 2012, Sharma & Giribet 2012, Sharma et al. 2011,
2012). We have only begun to scratch the surface as the true diversity and biogeography of Australasian Samoidae
is largely unknown. Detailed studies of the remaining taxa, as well as an examination of new material collected
from this region, will undoubtedly promote species discovery, and elucidate evolutionary histories and
biogeographic patterns.
Acknowledgements
We thank Peter Jäger and Julia Altmann (SMF) for the loan of Fijicolana tuberculata types and for their very kind
attention during the first author’s visit to the SMF. Images of Badessa ampycoides were kindly provided by
Guilherme Gainett (Instituto de Biociências, Universidade de São Paulo) and by Gonzalo Giribet (Museum of
Comparative Zoology, Harvard University). Carlos Víquez, Juan Mata and Randall Gutierrez (INBIO) provided
support to the first author for collecting in Costa Rica. Comments from Peter Schwendinger and Ricardo Pinto-da-
Rocha helped to greatly improve the manuscript. This study was funded by the Argentinean grants PICT 2011-
1007 from the Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT) and PIP 2012-0943 from the
Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET).
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