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On the taxonomic position of the enigmatic
genus Tonkinodentus Schileyko, 1992 (Chilopoda,
Scolopendromorpha): the first molecular data

Arkady A. Schileyko', Evgeniya N. Solovyeva'

| Zoological Museum of the Moscow Lomonosov State University, Bolshaya Nikitskaja Str. 6, Moscow,
103009, Russia

Corresponding author: Arkady A. Schileyko (schileyko1965@gmail.com)

Academic editor: Pavel Stoev | Received 3 February 2019 | Accepted 29 March 2019 | Published 17 April 2019
http://zoobank. org/F4D86F D0-41E8-47D7-855F-523506B9I921E

Citation: Schileyko AA, Solovyeva EN (2019) On the taxonomic position of the enigmatic genus Tonkinodentus
Schileyko, 1992 (Chilopoda, Scolopendromorpha): the first molecular data. ZooKeys 840: 133-155. https://doi.
org/10.3897/zookeys.840.33635

Abstract

The taxonomic position of the monotypic Vietnamese genus TJonkinodentus Schileyko, 1992 (for T
lestes Schileyko, 1992) has been considered in the light of the first obtained molecular data. Both mo-
lecular (28S rRNA) and morphological data support the position of this extraordinary eye-less genus
within the family Scolopendridae Leach, 1814, a sighted clade, and thus suggests the polyphyly of blind
scolopendromorphs. The species diagnosis has been amended and color images of T’ lestes provided for
the first time.

Keywords
Extended redescription, molecular analysis, Scolopendridae, taxonomic position, Tonkinodentus, 18S

rRNA, 28S rRNA

Introduction

The monotypic genus 7onkinodentus Schileyko, 1992, based on T. lestes Schileyko, 1992,
was described from Vietnam by a single adult specimen lacking the ultimate pair of legs.
This enigmatic taxon, originally collected from Boun Ma Thuout in Dak Lak Province
(Fig. 1), is extraordinary in lacking eyes (Figs 2, 4) but otherwise scolopendrid-like in

Copyright Arkady A. Schileyko, Evgeniya N. Solovyeva. This is an open access article distributed under the terms of the Creative Commons Attribution Li-
cense (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

134 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

all other aspects. According to Schileyko (1992), the presence of forcipular tooth-plates
and a trochanteroprefemoral process (Figs 3, 5, 6) and the absence of a sternal trans-
verse suture (Fig. 7) place Tonkinodentus in Theatopsinae Verhoeff, 1906 (in the sense
of Schileyko 1992). Shelley (1997) synonymised Theatopsinae with Plutoniuminae
(= Plutoniumidae) Bollman, 1893 and also removed Tonkinodentus from this subfamily.

In 1994 another (complete) subadult specimen of 7! /estes (Fig. 8) was found in
Dong Nai Province (Fig. 1), and the species was redescribed by Schileyko (2007). As a
result, it turned out that Zonkinodentus has (as the overwhelming majority of the sub-
family Scolopendrinae Leach, 1814) paired sternal longitudinal sutures (Fig. 7), slit-like
spiracles (Fig. 9) covered by “flap” (a synapomorphy that is unique for Scolopendrinae),

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Figures 1-4. | Map of Vietnam showing the places of collection (black circles) of the holotype (Dak
Lak Province) and the second specimen (Dong Nai Province) of Tonkinodentus lestes Schileyko, 1992;
Tonkinodentus lestes Schileyko, 1992, holotype (Rc 6358) 2 general view, dorsally 3 general view, ventrally
4 head plate and LBS 1, dorsal view.

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 135

a well-developed and spinulated coxopleural process (Figs 10, 11), and regular (“com-
mon” sensu Schileyko 2009) ultimate legs (Figs 8, 12). Thus, 7’ /estes is morphologically
much closer to the sighted Scolopendrinae rather than to the blind Plutoniumidae. The
subsequent cladistic analyses of Edgecombe and Giribet (2004), Vahtera et al. (2012a),
and others (see below), which were based on both molecular and morphologic data,
proposed the monophyly of the blind scolopendromorphs (i.e. Cryptopidae sensu At-
tems, 1930); however, they did not contain data on Tonkinodentus.

Taking into consideration all these facts, Schileyko (2007) supposed Tonkinodentus to
be the first blind representative of Scolopendridae. Schileyko (2007: 71) also wrote that a
discussion concerning taxonomic position of this genus will be published elsewhere and left
Tonkinodentus unassigned to any subfamily. Thus, the aim of this paper is to specify a taxo-
nomic position of this enigmatic genus using the first molecular data as presented below.

Material and methods

All the studied material is deposited in the Zoological Museum of Moscow Lomono-
sov State University (ZMMU). The work was carried out based on the two specimens
of Tonkinodentus lestes (Rc 6358, holotype; Rc 6555, non-type). Abbreviations used are
LBS = leg-bearing segment(s), col. = collector. Specimens were examined both wet and
dry under various angles of direct illumination; the photos were taken using a Canon
EF-S 60 macro lens mm mounted on Canon EOS 300 camera and DeltaPix Invenio-
8DII digital camera. Lewis et al. (2005) and Bonato et al. (2010) were followed for
standard terminology of centipede morphology.

A tissue sample of 7’ estes was taken from the 75% ethanol preserved specimen
(voucher number Rc 6555 in ZMMU, collected in 1994). To avoid contamination,
extraction and amplification of the DNA were carried out in the ZMMU Laboratory
of Historical DNA. This laboratory was specially designed for work with samples from
museum specimens, which potentially have their DNA degraded. No previous work
on fresh tissues had been performed in this laboratory (Kruskop et al. 2018; Lebedev
et al. 2018). DNA was extracted twice; for the first time, it was extracted and purified
using the QlAamp DNA Minikit (Qiagen), which included an overnight lysis step at
56 °C and longer incubation with EB-buffer (5 min) at the purification step. For the
second time, DNA was extracted using a non-destructive method (Gilbert et al. 2007)
with the following modifications: incubation at 55 °C was performed for 8 h, DNA
purification was done with Qiagen PCR purification kit.

We amplified a fragment of the 28S rRNA nuclear gene. The DNA was highly
degraded, so short fragments (100-200 bp) were obtained using the combination of
internal primers designed for this study (Appendix 1, 2). Primer pairs were developed
manually using Bioedit (Hall 1999) and an alignment of candidate centipede sequences
from GenBank. First DNA extraction was successfully amplified with 28S-endF/28S-
endR primer pairs, but another pair of primers (startF_2 and IntR_2) worked only on
the second DNA extraction.

136 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

Figures 5-9. Jonkinodentus lestes Schileyko, 1992; Holotype (Rc 6358) 5 anterior margin of forcipular
coxosternite, ventral view 6 head, forcipular segment and LBS 1-2, ventral view 7 LBS 12, ventral view;
non-type (Rc 6555) 8 general view, laterally; Holotype (Rc 6358) 9 left side of LBS 2-4, lateral view; (asp)
— accessory spines, (bs) — basal sutures of tooth-plates, (cl) — chitin-line, (dhp) — dorsal half of process of

trochanteroprefemur, (im) — pleural intersclerite membrane, (lmd) — longitudinal median depression, (ms)
— medial suture, (pr) — leg pretarsus, (ps) — paramedian suture, (pt) — process of trochanteroprefemur, (sp)
— slit-like spiracle of LBS 3, (ss) — short caudo-lateral suture, (tp) — tooth-plate, (tr) — tarsungula, (ts) — leg
tarsal spur, (t1) — leg tarsus 1, (t2) — leg tarsus 2, (vhp) — ventral half of process of trochanteroprefemur.

The PCR program for amplification of short fragments included an initial denaturation
at 95 °C for 3 min, 45 cycles of 95 °C for 30 s, annealing temperature (see Appendix 1) for
30 sand 72 °C for 30 s, and a final extension of 72 °C for 6 min. All stages of the extraction
process included a negative control run in parallel. PCR products were visualized on a 1%
agarose gel. PCR product was sequenced via Evrogen on ABI PRISM 3500xl sequencer. All
sequences were deposited in GenBank under the following accession number MK517656.

Additional sequences of 28S rRNA and 18S rRNA of various scolopendromorphs

(including the members of Scolopendrinae, i.e. potential close relatives of 7’ lestes) were

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 137

downloaded from GenBank (see Appendix 3). Craterostigmus tasmanianus Pocock, 1902,
a member of Craterostigmomorpha, was used as an outgroup. We did not increase the
length and variability of our alignment by adding mitochondrial DNA data available for
this set of taxa (excluding 7’ /estes) in GenBank because the Chilopoda mtDNA sequenc-
es are very variable, much more than their nuDNA ones and there is a high possibility
of saturation of mtDNA while comparing distant taxa and because the cases of mito-
chonrial introgression are rather common. We hold to an opinion that in such situations
combining both nuDNA and mtDNA data in one alignment can lead to errors and is
better avoided, especially when, as in our case, the DNA fragment of the target specimen
is short and represents only one type of DNA markers (either nuDNA, or mtDNA).

Sequences of 7” /estes were checked and put in contig using Seqman 5.06 (Bur-
land 1999). Than contig and GenBank sequences were aligned with Geneious 11.1.5
(http://www.geneious.com) using Geneious Alignment. Subsequently, the alignment
was checked and manually revised if necessary using BioEdit Sequence Alignment
Editor v. 7.1.3.0 (Hall 1999). Two alignments were prepared for the following phylo-
genetic analysis: sequences of 28S only and concatenated alignment of 28S + 18S. We
did not cut the sequences of the other taxa to match the length of the 7’ /estes fragment,
the length of 28S alignment was 1743 b.p. and 1913 b.p. for 18S alignment. 18S se-
quences were added to improve the resolution of the resulting trees. Genetic distances
were calculated using MEGA 6.1 (Tamura et al. 2013).

The optimum partitioning schemes were identified with PartitionFinder (Lanfear et
al. 2012) using the greedy search algorithm under the AIC criterion: GTR + 1+ G for 28S
and SIM + 1+ G for 18S. Phylogenetic trees were reconstructed under Bayesian criteria
(BI) and the maximum likelihood (ML). Bayesian inference (BI) was performed in Mr-
Bayes v. 3.1.2 (Ronquist and Huelsenbeck 2003) with two simultaneous runs, each with
four chains, for 8 million generations for 28S and 12 million generations for 28S + 18S.
We checked the convergence of the runs and that the effective sample sizes (ESS) were all
above 200 by exploring the likelihood plots using TRACER v. 1.5 (Rambaut and Drum-
mond 2007). The initial 10% of trees were discarded as burn-in. Confidence in tree
topology was assessed by posterior probability (PP) (Huelsenbeck and Ronquist 2001).

The ML trees were generated in IQ tree (Nguyen et al. 2015) using ultrafast boot-
strap = 10000 (UFBoot, Minh et al. 2013). A model for the 28S (GTR+F+I+G4)
alignment was selected using ModelFinder (Kalyaanamoorthy et al. 2017), and a par-
titioning model for the 28S + 18S alignment were calculated with IQtree (Chernomor
et al. 2016): GT R+F+I+G4 for 28S and TNe+I+G4 for 18S.

Results
1. Amended diagnosis and redescription of Tonkinodentus lestes Schileyko, 1992
Schileyko (2007) redescribed 7’ /estes in detail, but the black-and-white photographs

are far from satisfactory. Below we present a new diagnosis and description of this spe-
cies accompanied by color photographs.

138 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

Figures 10-14. Zonkinodentus lestes Schileyko, 1992; Holotype (Re 6358) 10 LBS 20-21, ventro-lateral
view I | LBS 20-21, ventral view; non-type (Rc 6555) 12 LBS 21 and ultimate legs, dorso-lateral view 13
head plate and LBS 1, dorsal view 14 LBS 13, ventral view; (as) — apical spine(s) of coxopleural process,
(cp) — coxopleural process, (eps) — coxopleural posterior spine, (cs) — corner spine of ultimate prefemur,
(f) — femur, (g) — gonopod, (pf) — prefemur, (ps) — paramedian sutures, (sas) — subapical spine(s) of coxo-
pleural process, (t) — tibia, (t1) — tarsus 1, (t2) — tarsus 2, (us) — ultimate sternite, (vlp) — distal ventro-

lateral process, (vs) — ventral spine(s) of coxopleural process.

Family Scolopendridae Leach, 1814
Subfamily Scolopendrinae Leach, 1814

Genus Jonkinodentus Schileyko, 1992

Type species. Zonkinodentus lestes Schileyko, 1992 (by monotypy).
Range. Central Vietnam, Dak Lak (Darlak) Province; South Vietnam, Dong Nai

Province.

On the taxonomic position of the enigmatic genus Tonkinodentus Schileyko, 1992... 139

Tonkinodentus lestes Schileyko, 1992
Figures 2-19

Tonkinodentus lestes Schileyko 1992: 13.
Tonkinodentus lestes: Schileyko 1995: 74.
Tonkinodentus lestes: Schileyko 2007: 83.

Locus typicus. Central Vietnam, Dak Lak (Darlak) Province, environs of Boun Ma Thuot.

Material. Dak Lak (Darlak) Province, ca 15 km of Buon Ma Thuot, Eakmat,
450 m, 1—5.05.1986, col. L.N. Medvedev, 1 spec. (holotype, Rc 6358); Dong Nai
Province, Ma Da Forest, Dipterocarpus area, soil samples, 19.10.1994, col. N.V. Beli-
aeva, 1 spec. (Rc 6555).

Diagnosis. Cephalic plate lacking any sutures, its posterior margin overlapped
by tergite 1; eyes absent (Figs 4, 13). Forcipular tooth-plates well developed and rela-
tively short, with 7 teeth arranged in 2 parallel rows in a chess-board pattern (Fig.
5); trochanteroprefemoral process bisected sagittally (Figs 5, 6). Sternites 2—20 with
paramedian sutures (Figs 7, 14). Pleuron with intersclerite membrane clearly visible;
spiracles triangular with a 3-part “flap”, slit-like entrance and deep atrium. 21 LBS;
the ultimate one visibly shorter than penultimate (Fig. 15). Leg with tarsus 1 consider-
ably longer than tarsus 2, with both tarsal spur and pretarsal accessory spines. Ultimate
sternite with poorly developed longitudinal median depression in caudal half. Cylin-
drical coxopleural process well developed, with spines (Figs 10, 16, 17). Ultimate legs
of “common” shape (sensu Schileyko 2009; Figs 8, 12); femur, tibia, and tarsus 1 each
with an apically rounded distal ventro-lateral process (Figs 8, 12, 18).

Composite redescription. [data concerning the non-type specimen in square
brackets]

Length of body ca 45 [34] mm. Color in ethanol: entire animal uniformly yellow-
brownish (Figs 2, 3) [pale yellow, nearly white; Fig. 8]. Body and legs with a very few
minute setae.

Antennae of 19 articles (in the both specimens left antenna of 19 and right one of
18, as the corresponding apical article seems to be broken off), reaching the anterior
margin of tergite 5 [5.5—6] when reflexed. Basal articles 6 or 7, with a very few long
setae, subsequent articles densely pilose. Basal antennal articles flattened.

Cephalic plate (Figs 4, 13) without any sutures, rounded and remarkably narrower
than tergite 1; its posterior margin covered by the latter. No light spots at the place of ocelli.

Maxillae 2: the second article of telopodite distally with dorsal spur. Dorsal brush
very poorly visible, consisting of short, delicate and transparent setae; apical setae no
longer than pretarsus. Uniformly brown pretarsus (Fig. 19) simple (not pectinate) and
claw-shaped, as long as 1/3—1/4 of the length of the apical article of telopodite; pretar-
sus with 2 thin accessory spines.

Forcipular segment: coxosternite with shortly branched medial suture which is as
long as 1/3 of coxosternal length; 2 short sutures stretched caudo-laterad from median

140 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

diastema (Fig. 5) [all coxosternal sutures very hardly visible] in the form of an angle of
ca 60° [ca 70°]; chitin-lines short but well developed (Fig. 6). Tooth-plates definitely
wider than long [visibly higher than in the holotype]; height of tooth margin increas-
ing medially. Each tooth-plate with 7 teeth, fused to various degrees and arranged in 2
parallel rows in a chess-board pattern (Fig. 5), the lateral tooth is the shortest and the
most isolated. Basal sutures of tooth-plates form a nearly straight line. Trochantero-
prefemural process well developed, divided sagittally into 2 (dorsal and ventral) halves
(Figs 5, 6), each half with 2 or 3 lateral tubercles [dorsal halves of both processes with
3, ventral ones (which are visibly smaller) with 2]; the apical end of this process is con-
siderably higher than corresponding tooth-plate. Tarsungula (Fig. 6) of normal length
(left one broken off apically in the holotype), ventrally with 2 blunt ridges.

Tergite 1 without sutures (Figs 4, 13), tergite 2 with visible incomplete paramedian
sutures, tergites 3—20 with well-developed and complete ones (Fig. 15), tergite 21 with
complete median suture. Tergites 15/16—20 with poorly developed lateral margination
posteriorly; only tergite 21 definitely marginate. Tergite 21 nearly as wide as long and
not narrowed caudad (Fig. 15); its lateral sides slightly rounded and posterior margin
evidently rounded. Tergites lacking any median keel.

Sternites 2—20 (Figs 7, 14) with lateral sides practically parallel; 2-20 with com-
plete paramedian sutures; sternites 6/7—18/19 with a well-developed longitudinal me-
dian depression, which is wide and deep [very shallow]. Ultimate sternite long and
very narrow (Fig. 11), at least twice as long as wide at base [1.5:1; Fig. 17], very slightly
narrowing caudad; its posterior margin practically straight [with strongly rounded cor-
ners]. Endosternites not recognizable.

Composition of pleuron (Fig. 9) usual for Scolopendrinae, intersclerite membrane
well visible. Elongated spiracles triangular with a typical for this subfamily 3-valved
“flap” which covers slit-like entrance in a well-developed atrium (Fig. 9).

Legs (Figs 7, 9) with tarsus 1 considerably longer than tarsus 2, legs 1-18 [1-19]
with tarsal spur (legs 19-21 of the holotype are lost). Pretarsus long (approximately as
long as %4 of tarsus 2), legs 1-20 with well-developed accessory spines.

Ultimate LBS visibly shorter than penultimate (Fig. 15). Coxopleuron (excluding
coxopleural process) visibly longer than sternite 21 (Figs 11, 16), its coxal part very densely
pierced with coxal pores of various size, only this coxopleural process and a narrow posteri-
or area remaining poreless [this posterior area visibly broader than in the holotype]. Short,
cylindrical coxopleural process (Figs 10, 16) slightly curved dorsad [definitely curved me-
dially and very slightly dorsad], with 2 apical, 2 subapical, and 2 ventral spines close to
its base [with 3 apical, 1 subapical, and 1 lateral spine]; 1 or 2 [1] spines at posterior
margin of coxopleuron. Coxopleural process practically reaches the caudal margin of the
ultimate tergite. Caudal margin of ultimate pleuron virtually straight and lacking spines;
coxopleural surface with scattered minute setae. Gonopods well developed (Figs 11, 17).

[Ultimate legs (Figs 8, 12) ca 7 mm long, relatively slender (width of prefemur ca
0.7 mm), prefemur definitely flattened dorsally, other articles cylindrical. Prefemur, fe-
mur and tibia practically of the same length (ca 1.7 mm), tarsus 1 considerably longer

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 141

17 j
Figures 15-20. Zonkinodentus lestes Schileyko, 1992; Holotype (Rc 6358) 15 LBS 20-21, dorsal view;
non-type (Rc 6555) 16 LBS 21 and ultimate prefemora, ventro-lateral view 17 LBS 21 and right ultimate

prefemur, ventral view 18 femur, tibia, and tarsus 1 of left (lateral view) and right (medial view) ultimate
leg; Holotype (Rc 6358) 19 maxillae 2 and anterior part of forcipular segment, ventral view; Cormo-
cephalus dentipes Pocock, 1891, adult Rc 7013 20 anterior margin of forcipular coxosternite, ventral view;
(as) — apical spines of coxopleural process, (asp) — accessory spines, (at) — apical article of telopodite of
maxilla 2, (ep) — coxopleural process, (eps) — coxopleural posterior spine, (cs) — corner spine of ultimate
prefemur, (dhp) — dorsal half of process of trochanteroprefemur, (f) — femur, (g) — gonopod, (Im) — lateral
margination, (Is) — lateral spine(s) of coxopleural process, (mds) — median suture, (ms) — medial spine(s)
of ultimate prefemur, (pa) — coxopleural posterior poreless area, (pr) — pretarsus, (ps) — paramedian
suture(s), (sas) — subapical spine(s) of coxopleural process, (t) — tibia, (tl) — tarsus 1, (us) — ultimate
sternite, (ut) — ultimate tergite, (vhp) — ventral half of process of trochanteroprefemur, (vlp) — distal
ventro-lateral process, (vls) — ventrolateral spine(s) of ultimate prefemur, (vms) — ventromedial spine(s)
of ultimate prefemur.

142 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

than tarsus 2 (Figs 8, 12, 18), the latter twice as long as pretarsus. Ventral surface of
prefemur spineless, left prefemur with 23 and right one with 20 small ventrolateral
spines, remaining ones (23 on left prefemur, 22 on right) disposed ventromedially,
medially and dorsomedially (Figs 12, 16, 17). The spines grouped in very indistinct
rows or scattered chaotically; corner spine well developed (Figs 12, 16), with 2 apical
spines. No tarsal spur; pretarsus slender, sharply contrasting to much thicker tarsus 2,
accessory spines absent. Tarsus 1 and tibia visibly broadened apically; femur, tibia and
tarsus 1 each with a characteristic distal process ventro-laterally, the latter short and
rounded apically (Fig. 8, 12, 18)].

Remarks. The known material consists of two specimens only, neither of which are
in perfect condition. More material is needed to investigate the anatomy (e.g. peristo-
matic structures, foregut, gizzard).

All differences between the holotype and the second specimen are explicable by the
latter being a subadult. The much paler and considerably softer cuticle the second spec-
imen suggests that it is newly moulted. Because of this, some delicate structures (e.g.
forcipular sutures, leg spurs) are less evident than in the holotype. The most delicate
parts (maxillae, antennae, legs) are somewhat deformed (wrinkled) in the holotype,
but in the second specimen, the ventral surfaces of the apical articles of the ultimate
legs are deformed (unnaturally concave).

Eight specimens of Cormocephalus dentipes Pocock, 1891 (Re 7518, 7013, 7028,
7231, 7233) from India (Assam and Punjab states), Western Nepal and Indonesia
(Sumatra, Medan) demonstrate virtually the same structure of the sagittaly bisected
process of the forcipular trochanteroprefemur (Fig. 20). As for the chess-board pattern
of the arrangement of the teeth of the forcipular tooth-plates in Zonkinodentus (Fig. 5),
it is unique among the Scolopendromorpha.

Discussion. ‘The genus Jonkinodentus conforms to the Scolopendrinae and differs
from members of both the Plutoniumidae and Cryptopidae Kohlrausch, 1881 by: (1)
the presence of paired sternal longitudinal sutures (Figs 7, 14) vs single median suture,
(2) the slit-like spiracles are covered by a “flap” (a synapomorphy that is unique for
Scolopendrinae; Fig. 9), with the longitudinal axis of the spiracle parallel to the such of
the body vs open oval spiracles, (3) the well-developed, spinulated coxopleural process
(Figs 10, 16) vs its virtual absence, and (4) the ultimate legs of “common” shape (sensu
Schileyko 2009; Figs 8, 12) vs enlarged, “pincer-shaped” ones in Plutoniumidae or
“pocket knife-shaped” ones in Cryptopidae. Tonkinodentus also sharply differs from the
typical cryptopids (= Cryptops Leach, 1814) by having: (1) well-developed forcipular
tooth-plates with strongly chitinized teeth, (2) a forcipular trochanteroprefemur with
a well-developed process, (3) sternites without transversal sutures, and (4) prefemur of
ultimate legs with numerous spines (Fig. 17).

Summing up, the genus 7onkinodentus is morphologically the typical representa-
tive of the subfamily Scolopendrinae (and namely of the former tribe Scolopendrini
Leach, 1814) and is the most similar to the genus Scolopendra L., 1758, but differs read-
ily from the latter by the absence of eyes and the peculiarities of the forcipular segment.

On the taxonomic position of the enigmatic genus Tonkinodentus Schileyko, 1992... 143

2. Results of the molecular analysis

WANS Sequence characteristics

We obtained 175 b.p. of 28S rRNA of Tonkinodentus lestes. The complete matrix in-
cluded sequences from 40 species. Information on the length of 28S and 18S frag-
ments and variability is given in Appendix 4 (all data shown for ingroup only). Uncor-
rected mtDNA genetic distances are given in Appendix 5, 6 (below diagonal).

2.2. Phylogenetic analysis

The results of the phylogenetic analysis are presented in Figures 21 and 22. BI and ML
analyses yielded trees that demonstrated essentially similar topologies. Trees based on
28S and 28S + 18S alignments are also rather congruent. Relations of blind species are
not resolved in the 28S phylogenetic tree, but non-blind group represents a clade (PP =
0.91, BS = 92). The monophyly of both Scolopendrinae (PP = 0.95, BS = 95) and Oto-
stigminae (PP = 0.94, BS = 100) was supported (Fig. 22). The 28S + 18S phylogenetic
tree shows lesser values of support (PP = 0.86, BS = 82) for the non-blind clade, but
relations within the blind group are better resolved: two species of Theatops Newport,
1844 form a clade (PP = 0.66 (not shown), BS = 97), Newportia Gervais, 1847 and
Scolopocryptops Newport, 1844 form another clade (PP = 0.098, BS = 98), and species
of Cryptops Leach, 1814 form the third clade (PP = 1, BS = 100). According to both
28S and 28S + 18S topologies, blind Tonkinodentus lestes is included in the non-blind
clade (= Scolopendridae).

Theatops erythrocephalus
heatops posticus

Newportia quadrimeropus

ewportia longitarsis
Scolopocryptops sexspinosus
Scolopocryptops miersii

Cryptops spinipes

Cryptops weberi

Cryptops australis

Newportia monticola

ryptops trisulcatus

0.96/95 Campylostigmus decipiens
-/100 Campylostigmus orientalis
. Cormocephalus aurantiipes
Cormocephalus monteithi
—— Tonkinodentus lestes ZMMU S-6555

blind

lopendra marginata

Notiasemus glauverti

Akymnopellis chilensis

Digitipes cf. barnabasi 1

Rhysida afra

Otostigmus astenus
Otostigmus caraibicus

Digitipes sp. 1
o.99/994 Digitipes cf. coonoorensis
t-)974 Digitipes cf. barnabasi 2

Otostigmus rugulosus
Edentistoma octosulcatum
Alipes
Ethmostigmus rubripes
Rhysida nuda
0.91/- Sterropristes violaceus

Craterostigmus_tasmanianus

ocellate

21

0.3

Figure 21. Phylogenetic BI tree reconstructed from alignment of the nuclear gene 28S. (Numbers on tree
nodes indicate posterior probabilities (PP > 90) and bootstrap values (BS > 75) for BI/ML, respectively).

144 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

197 Theatops erythrocephalus
Theatops posticus
0.99/100 Newportia quadrimeropus
0.98/98 Newportia longitarsis
: 0.90/99 Newportia monticola
Scolopocryptops sexspinosus
1/98 Scolopocryptops miersii
4I- Cryptops spinipes
1/99 Cryptops australis
4/100 Cryptops weberi
0.96/88 Cryptops trisulcatus
0.98/99 Campylostigmus decipiens
Campylostigmus orientalis
Cormocephalus aurantiipes
Notiasemus glauerti
Cormocephalus monteithi
Tonkinodentus lestes ZMMU $-6555
/9 Scolopendra cingulata
0.45/95 Scolopendra subspinipes
Scolopendra morsitans
Asanada socotrana
1/98 Asanada brevicornis
-/98 Scolopendra viridis
189 Arthrorhabdus formosus
Hemiscolopendra marginata
Akymnopellis chilensis
182 Alipes grandidieri
Otostigmus astenus
Edentistoma octosulcatum

ooeper Digitipes sp.

blind

ocellate

Digitipes cf. coonoorensis
0.947700;/24 Digitipes cf. barnabasi 2
4 ‘178— Ethmostigmus rubripes
Lt Rhysida nuda
0.96/- Sterropristes violaceus
Otostigmus caraibicus
Otostigmus rugulosus
Digitipes cf. barnabasi 1
-176 Rhysida afra
Craterostigmus tasmanianus

22

0.02

Figure 22. Phylogenetic BI tree reconstructed from concatenated alignment of the nuclear genes 28S +
18S. (Numbers on tree nodes indicate posterior probabilities (PP > 90) and bootstrap values (BS > 75)
for BI/ML, respectively).

3. The taxonomic position of Zonkinodentus and the problem of mono- vs para-

phyly of the blind scolopendromorphs

The question of the correct taxonomic position of Tonkinodentus (in fact, the first eye-less
scolopendrid) is connected directly with the problem of mono- vs paraphyly of the blind
scolopendromorphs. An origin of the family Cryptopidae sensu Attems (1930), or the
“blind clade” sensu Vahtera et al. (2012a), which includes all three eye-less scolopendro-
morph families (Cryptopidae, Plutoniumidae, and Scolopocryptopidae Pocock, 1896) is
a matter of a long discussion. Schileyko (1992) argued the monophyly of the blind scolo-
pendromorphs, stating that the group “Cryptopidae” is not a natural taxon, and tried to
support this by producing the first character matrix for the order (Schileyko 1996). ‘This
was, however, quite limited and included only 15 genera and eight characters. ‘This view-
point was, in part, supported by Shelley (1997: 106), who wrote: “... no longer should
the present division [of order Scolopendromorpha], based primarily on the presence or ab-
sence of eyes, be uncritically accepted”. Shelley (2002: 2) later wrote: “Based on anatomi-
cal and biogeographical considerations (discussed by Shelley (1997)), I return the Scolo-
pocryptopinae to full family status from a subfamilial position under the Cryptopidae.”
It is interesting that the results of the purely morphological investigations demon-
strate that the para- vs monophyly of the blind scolopendromorphs depends on the
parameters of the analysis (Edgecombe and Koch 2008; Edgecombe and Koch 2009;
Di et al. 2010) and that “The status of blind Scolopendromorpha as a grade or clade
remains an open question” (Edgecombe and Koch 2009: 311). These conclusions have

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 145

been supported by Koch et al. (2010: 70): “The shortest cladograms include two alter-
native resolutions of blind scolopendromorphs”.

In contrast, both molecular and/or combined analyses supported the monophyly
of the eye-less clade (Edgecombe and Giribet 2004; Vahtera et. al. 2012a). Edgecombe
and Giribet (2004: 125) wrote: “The cryptopid clade is present across most of param-
eter space for combined morphological and molecular data (...), leading us to favor
the hypothesis that loss of ocelli in Cryptopidae occurred once and defines a mono-
phyletic group”. Also Vahtera et. al. (2012a, 2013) considered a single loss of ocelli in
Scolopendromorpha as the most parsimonious. Confirming these conclusions Bonato
et al. (2017: 2) stated that Plutoniumidae, Cryptopidae and Scolopocryptopidae are a
“... well-supported monophyletic subgroup, informally labelled as the “blind clade’...”.

Morphology and Sanger sequence data reviewed above have been inconclusive
with regards to the monophyly of a clade uniting the blind scolopendromorphs (except
for Tonkinodentus); this grouping is robustly supported by phylogenomic data (Fernan-
dez et al. 2016). All of 20 analyses using different gene partitions, optimality criteria
(Bayseian Inference or Maximum Likelihood), or tree-inference algorithms recover
this group with strong support.

The only mention of Tonkinodentus within this discussion has been made by Vahtera
et al. (2012a: 14), who wrote: “Our data are lacking the monotypic blind scolopendrid
genus Tonkinodentus Schileyko, 1992. Morphology supports an assignment of this rare
genus to Scolopendridae (Schileyko 2007) but this hypothesis remains yet to be tested in
terms of the molecular data. As such, although we postulate a single origin for blindness
in three families of Scolopendromorpha, an independent loss of ocelli within Scolopen-
dridae (in Zonkinodentus) is probably based on published morphological evidence for
the affinities of Tonkinodentus’. Summing up, the results of the first molecular approach
applied to this peculiar genus should be of the special importance for this discussion.

As it was already noted above, Schileyko (2007) assigned Tonkinodentus to the
family Scolopendridae (sensu lato), so the precise taxonomic position of Tonkinoden-
tus within the family remains indefinite. In the most current general review of scolo-
pendromorph genera, Edgecombe and Bonato (2011: 400) included this genus in
the former tribe Scolopendrini Leach, 1814, but provided no arguments for doing
so. Later, using a combined morphological and molecular approach, Vahtera et al.
(2013: 578) showed that “the tribe Asanadini [Verhoeff, 1907] nests within Scolo-
pendrini for molecular and combined datasets”, thus reducing both tribes, but with-
out formalizing their new statuses. The molecular data confirms that 7onkinodentus
nests in the family Scolopendridae, or in the subfamily Scolopendrinae (Figs 21,
22), and thus, the discovery of the first eye-less scolopendrid is confirmed.

Conclusions

Work with ancient DNA from long-preserved museum collections is now an impor-
tant and developing, but complicated, phylogenetic approach. In this study of 7. lestes,

146 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

DNA was so degraded and in so small an amount that two different methods of DNA
extraction were used, and only two short fragments of 28S rRNA were obtained.
Both morphological and the first molecular data unequivocally support the position
of blind Yonkinodentus inside sighted Scolopendridae. ‘The position of Tonkinodentus
among the members of Scolopendrinae (i.e. non-blind scolopendromorphs with slit-like
spiracles covered by a “flap”) is well confirmed by morphological data, but has quite
low nodal support in our phylogenetic analysis. More fresh materials are necessary to
complete both internal anatomical and molecular studies of this enigmatic scolopendrid.
The position of 7’ /estes within the sighted family Scolopendridae coincides with
hypothesis that blind scolopendromorphs are non-monophyletic, although phylog-
enomics strongly supports monophyly of a clade of the three obligately blind families.

Acknowledgements

We thank both collectors of the specimens of Tonkinodentus lestes. We are very grateful
to the Director of ZMMU Dr Mikhail Kalyakin for financial support of publication
payments and to Dr Vladimir Lebedev (ZMMU) for his consultations. Our sincer-
est thanks are to Prof. Pavel Stoev for his kind help in editing earlier versions of the
manuscript and to Dr Varpu Vahtera and Dr Gregory Edgecombe for their valuable
comments and suggestions for improvement during the review process. Our work has
been carried out within the state program “Taxonomic and chorological analysis of
the animal world, as a ground for study and conservation of the biological diversity”

(AAAA-A16-116021660077-3) of the Moscow Lomonosov State University.

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Appendix |

Primer pairs used in this study.

Fragment Primer name Sequence of 3'—5' Annealing temperature
1 28S-endF 3'-GGAGTCCCGGGAAGAGTTGTC-5' 56°C

1 28S-endR 3'-TACGGTCCGGCGCGAAAATCA-5'

D, startF_2 3'-CCGAGCGACCGAAAGGGAATC-5' 58 °C

2 IntR_2 3'-AGTCCGTCCCT TACAAAGAAAAGACAACTCT-5'

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 149

Appendix 2

Start_F2 28S-endF

Simplified scheme of the primer positions on 28S gene.

Appendix 3

Sequences used in this study.

species 28S 18S
Akymnopellis chilensis HQ402521 HQ402503
Alipes grandidieri HM453273 KF676422
Arthrorhabdus formosus HQ402522 HQ402504
Asanada brevicornis HQ402523 HQ402505
Asanada socotrana HQ402524 HQ402506
Campylostigmus decipiens HQ402525 HQ402507
Campylostigmus orientalis HQ402526 HQ402508
Cormocephalus aurantiipes HQ402527 HQ402509
Cormocephalus monteithi HM453274 AF173249
Craterostigmus tasmanianus HM453266 AF000774
Cryptops australis AY288708 AY288692
Cryptops spinipes AY288709 AY288693
Cryptops trisulcatus AF000783 AF000775
Cryptops weberi HQ402535 HQ402518
Digitipes cf. barnabasi | JN003983 =
Digitipes cf. barnabasi 2 JN003987 -
Digitipes cf. coonoorensis JN003979 =
Digitipes sp. JN003980 -
Edentistoma octosulcatum KM492928 KM492930
Ethmostigmus rubripes HM453276 KF676424
Hemiscolopendra marginata HQ402530 HQ402513
Newportia longitarsis HM453281 HM453236
Newportia monticola HQ402531 HQ402514
Newportia quadrimeropus HQ402529 HQ402511
Notiasemus glauerti KF676405 KF676456
Otostigmus astenus HQ402532 HQ402515
Otostigmus caraibicus HQ402533 HQ402516
Otostigmus rugulosus HQ402534 HQ402517
Rhysida afra HQ402536 -
Rhysida nuda HM453277 AF173252
Scolopendra cingulata HM453275 U29493
Scolopendra morsitans HQ402537 HQ402519
Scolopendra subspinipes HQ402538 HQ402520
Scolopendra viridis DQ222134 DQ201419
Scolopocryptops miersii HQ402528 JX422720
Scolopocryptops sexspinosus AY288710 AY288694
Sterropristes violaceus KF676377 KF676428
Theatops erythrocephalus HM453279 AF000776
Theatops posticus HM453280 AY288695
Tonkinodentus lestes MK517656 -

150 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

Appendix 4

Sequence characteristics. Cons. = conservative sites, Var. = variative sites, Pars.-Inf. = parsimony informa-

tive sites.
Nucleotide frequencies (%)
Locus Length (b.p.) Cons. Var. Pars.-Inf. TU C 7 G
28S 1743 945 744 381 18.5 Z3eF 23.8 31.9
18S 1913. 1662 237 120 23.4 24.7 23./. 28.2:
Appendix 5

Uncorrected p-distances (%) for sequences of 28S nuDNA gene for species (above diagonal). Standard

error estimates are shown above the diagonal.

i rE
Akymnopellis 124
1 | chilensis ;
2 |Alipes grandidieri 8.34 1.01
Arthrorhabdus 8.33 1.39
3 | formosus
4 |Asanada brevicornis | 5.07 7.00
5 |Asanada socotrana | 6.76 1.41
COURS | |i5.0% 7.00
6 | decipiens
COIR IE | acy 7.00
7 | orientalis
Cormocephalus 5.14 7.00
8 | aurantiipes
orm ceD are 5.14 1.96 | 0.88 | 1.91 | 1.41
9 | monteithi
10 | Cryptops australis | 17.96 1.77 f 2.05
11 | Cryptops spinipes 16.27 2.11
12 | Cryptops trisulcatus | 19.32 20.33} nic 2.18
13 | Cryptops weberi 11.26 12.79 | 7.90 1.66
Digitipes cf.
14 | barnabasi 1 Lae 1.27
Digitipes cf.
15 | barnabasi 2 10.58 14.62 | 34.04
16 | Digitipes sp. 1 10.89 1.01
Dene 10.44 11.40} 21.55| 20.67 | 22.12 0.00
17 | coonoorensis
ne 13.53] 8. 14,66 | 20.65 | 20.53 | 22.19 | 18.86] 11.28] 6.67
18 | octosulcatum
Ethmostigmus
i 8.71 | 4.61 | 9.29 | 4.39 | 7.71 | 4.96 | 6.40 | 6.19 | 7.49 | 20.47 | 21.47] 21.61) 12.76] 9.65 | 6.82
19 | rubripes
deep OpenaLe™ \ tears: | 6.08 | 5.95 nic | 9.32 | nlc |31.91
20 | marginata
ied ain 19.24 20.02 | 17.20] 15.80 | 17.70 | 19.25 | 18.47] 15.79 | 27.74 | 26.52 | 25.81 | 19.45 | 23.93 | 20.53
21 | longitarsis
22 | Newportia monticola| 8.42 | 8.04 | 11.03] 6.50 | 9.13 | 7.50 |10.11] 9.11 | 4.30 31.91
Newport 3.89 | 4.59 | 4.24 | 4.24 | 6.03 | 4.95 | 4.95 | 4.95 | 3.18 | nlc | n/c | nlc | 4.26] nlc | nic
23 | quadrimeropus
24 | Notiasemus glauerti | 11.24) 13.47|13.39] n/c |17.71] n/c | nlc | nie | 14.00} 20.17 | 20.45 | 21.51 | 22.26 | 15.90] 14.46

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992...

11 12 13 14

151

15

25 | Otostigmus astenus | 7.09 4.73 | 7.50 7.46 | 7.08 |20.41| 20.53] 21.55 | 12.55| 8.50 | 6.79
Orosigunts 7.63 | 4.78 9.04 | 7.03
26 | caraibicus
27 | Otostigmus rugulosus| 7.74 | 4.95 8.75 | 6.55
Scolopendra
28 | cingulata 8.25 14.33 | 16.05
penlopen aie 5.57 | 9.05 12.15 | 14.75
29 | subspinipes
30 | Scolopendra viridis | 7.10 | 9.75 14.68 | 14.40
pe Milo 15.59| 18.52 nic |46.33
31 | miersti
Senlepodyaioes 20.70 | 25.28 18.90 | 22.38 25.53| 25.82
32 | sexspinosus
Sanlonenara 4.36 | 6.55 nic [34.04
33 | morsitans
Seerroprastes 12.35| 6.15 | 13.83] n/c [13.15] nlc | n/c | nlc [14.24] 21.64] 21.22| 22.70] 20.85 | 10.65] 7.56
34 | violaceus
35 [Rhysida aia | 8.09 13 | 687 | 748 [798 5102
36 | Rhysida nuda 8.96 4.57 | 7.14 | 4.76 | 7.62 | 6.84 | 7.63 9.68 | 7.23
tDeataps 10.68 | 11.07 16.08 | 20.00| 11.77| 16.41 | 15.40
37 | erythrocephalus
38 | Theatops posticus 8.63 | 8.69 n/c | 20.57
Tonkinodentus lestes
39 |ZMMU S$-6555 2.29 | 2.29 n/c | nic

Appendix 5 (continuation)

Uncorrected p-distances (%) for sequences of 28S nuDNA gene for species (above diagonal). Standard

error estimates are shown above the diagonal.

B 29 | 30

0.68 | 0.84 | 0.60 | 0.65

0.66 | 0.89 | 0.84 | 0.78

0.72 | 0.84 | 0.69 | 0.70

0.84 | 0.66 0.68 | 0.73 | 0.73 0.72

Asanada socotrana 1.72 | 1.96 0.69 | 0.79 | 0.63 | 0.67

Campylostigmus 0.75 | 0.86 | 0.70 | 0.79
decipiens

yells Spi Saeed 0.80 | 0.95 | 0.75 | 0.81
orientalis

sibs Soret 0.87 | 0.87 | 0.65 | 0.77
aurantipes

Cormocephalus 0.69
monteithi

Cryptops australis 2.00

Cryptops spinipes 1.97

1.94

0.98 | 0.92 0.82 | 0.79 | 0.79 0.82

2.27 | nic 2.08 | 1.84 | 1.92

1.55 | 6.81 1.50 | 1.42 | 1.38

Digitipes sp. 1 1.36 | 0.89 1.90 | 0.92 | 0.84 | 0.84 1.44

152 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

7 20 [a | 2 [2

1.25

Edentistoma 1.46 | 1.44

octosulcatum

0
Al

Ethmostigmus 0.66
Hemiscolopendra

‘ 0.91
marginata

NEUE ont e 17.49| 26.09] 16.61] 18.10 00 | 1.08

1.12
monticola

Newportia
quadrimeropus

Otostigmus
caraibicus
Otostigmus ‘ : 4.71 | 3.87 0.86 | 0.68 | 0.65
rugulosus

Scolopendra 12.88| 14.71 : ; 10.67 | 10.50 | 10.69 0.51
cingulata

subspinipes

Scolopendra viridis |12.92| 10.91] 12.83 ; 8.77 | 9.67 | 7.68

a cot eg : 16.53] 16.63 | 15.90] 20.96] 17.11] 16.16
miersii

Olaparryp sips 25.00| 25.21 24.44|24.44|24.44|21.70|21.19| 19.48
sexspinosus

Scolopendra ; 6.42 | 4.24 | 2.18 | 5.00
morsitans

Sterropristes

; 7.49 | 7.52 384] 608] n/c /26.32] ale | nie ]15.06 7.56 | 6.61 | 7.44 12.54
violaceus

Rhysida afra 8.16 | 4.48 | 8.26 | 6.78 | 6.62 [20.31] 9.43 | 3.90 |13.51| 5.64 | 6.31 | 5.46 [11.46] 9.38 | 9.22
Rhysida nuda 5.73 | 3.54 | 8.45 | 4.20 | 8.09 [16.53] 11.79| 4.24 |16.24] 6.67 9.84

Saateps 11.75] 15.88 11.43} 11.91} 11.21] 11.36} 10.49} 11.11
erythrocephalus

Theatops posticus 10.17} 12.50} 11.28] 10.76
Tonkinodentus

lestes ZMMU : 2.86 | 6.94 | 2.29 | 2.86
S-6555

Appendix 5 (continuation)

Uncorrected p-distances (%) for sequences of 28S nuDNA gene for species (above diagonal). Standard
error estimates are shown above the diagonal.

31 37_| 38 | 39
1 |Akymnopellis chilensis 1.06 O75. |° 165 0.96 | 1.03
2 |Alipes grandidieri 1.40 0.99 | 1.27 103% [i203
3 | Arthrorhabdus formosus 1.07 0.79 | 1.64 1.00 | 1.28
bn |Asoneda brewers 0.86 0.89 [1.32
5 | Asanada socotrana 1.07 0.81 | 1.92 0.93 | 1.82
6 | Campylostigmus decipiens 0.95 n/c 0.71 n/c 0.77 0.96 1.16 1.00 1.67
7 | Campylostigmus orientalis 0.95 n/c 0.77 n/c 0.80 1.07 1.20 1.09 1.67
8 | Cormocephalus aurantiipes 0.99 n/c 0.68 n/c 0.87 1.03 1.19 1.10 1.60

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 153

31 37_| 38 | 39

9 | Cormocephalus monteithi 1.22 0.67 1.72 0.85 1.04
10 | Cryptops australis n/c
11 | Cryptops spinipes n/c
12 | Cryptops trisulcatus n/c
13. | Cryptops weberi 1.65
14 | Digitipes cf. barnabasi 1 n/c
15 | Digitipes cf. barnabasi 2 n/c
16 | Digitipes sp. 1 n/c
17 | Digitipes cf. coonoorensis ; . : . . n/c
18 | Edentistoma octosulcatum n/c 1.38 n/c
19 | Ethmostigmus rubripes 0.95 0. 78 0.93 1.25 1.17
20 | Hemiscolopendra marginata 0.92 iors 0.70 n/c 0.91 0.88
21 | Newportia longitarsis 1.34 1.20 1.24
22 | Newportia monticola 0.93 imeierd 0. 89 n/c 1.03 1.24
23 | Newportia quadrimeropus 0.97 peels | Ae ie seve aa Bled 1.12 1.30
24 | Notiasemus glauerti n/c n/c n/c
25 | Otostigmus astenus 1.08 0. 7 1.40 097° |. 1.17
26 | Otostigmus caraibicus 1.06 1.18
27 | Otostigmus rugulosus 1.07 1.17
28 | Scolopendra cingulata 1.37 1.87
29 | Scolopendra subspinipes 1.09 1.06
30 | Scolopendra viridis 1.08 1.13
31 | Scolopocryptops miersii 1.42
32 | Scolopocryptops sexspinosus n/c n/c
33. | Scolopendra morsitans 8. _ : ‘ : . 1.29
34 | Sterropristes violaceus sor [we [ae Tas n/c n/c
35. | Rhyida afi 688 ior | 1.04
36 | Rhysida nuda 18.56 605 [7.24 0.95 | 1.17
37 | Theatops erythrocephalus 18.53 rio.o1 | 752 | 15.99 [iiss [ire | 0.82 1.41
Sa [Ieee i909 | we [37 | we [ios | om | 7m [134
39 | Tonkinodentus lestes ZMMU S-6555 4.00 | nic | 3.43 | nlc | 2.29 | 2.86 | 4.00 | 3.43

Appendix 6

Uncorrected p-distances (%) for sequences of 18S nuDNA gene for species (above diagonal). Standard

error estimates are shown above the diagonal.

Be

Pe romeenclss 0.41 | 0.37 | 0.32 | 0.38 | 0.42 | 0.30 | 0.40 | 0.47 | 0.42 | 0.43 | 0.30 | 0.29 | 0.37 | 0.36 | 0.28 | 0.45
chilensis

2 |Alipes grandidieri | 0.42) 0.41 | 0.31 | 0.36 | 0.45 | 0.46 | 0.43 | 0.45 | 0.36 | 0.35 | 0.28 | 0.26 | 0.33 | 0.42

3 2.86 | 3.01 0.33 | 0.42 | 0.32 | 0.31 | 0.45 | 0.49 | 0.45 | 0.46 | 0.32 | 0.28 | 0.38 | 0.33 | 0.30 | 0.47

4 |Asanada brevicornis | 1.90 | 1.95 | 2.23 0.28 | 0. 5 : ’ 0.44 | 0.44 | 0.24 | 0.21 | 0.31 | 0.28 | 0.21 | 0.41

5 0.31 | 0.33 | 0.45 | 0.53 | 0.48 | 0.47 | 0.34 | 0.32 | 0.36 | 0.34 | 0.30 | 0.48

6 | Campylostigmus |, 351 1 94] 1.49] 0.74] 1.24 0.44] 0.44} 0.14] 0.19 | 0.30 | 0.29 | 0.25 | 0.33
decipiens

PNAS e athe 1.91] 2.17 | 2.08} 1.06 | 1.99 0.42 | 0.42 | 0.10 | 0.18 | 0.33 | 0.29 | 0.20 | 0.43

orientalis

8 3.33 | 3.80 | 2.32]3.03] | 0.34] 0.32] 0.39 0.43 | 0.38 | 0.49
0.47 [0.44 [053
10 | Grypeopserisuleatus [3.89 3.65 |402[3.17[3.81[241[3.04|1.99|2.72| [0.34 0.43 [0.44 [0.43 |0.41 [0.40 0.47
11 | Cryptops weberi _| 3,93 [4.02 | 4.06 4.02 | 4.06| 4.06 r3.50| 4.09 [2.65] 3.25]2.65|2.98|1.94| 0.43/04 F043 [0.42 | 0.42 0.42 [0.43 0.40 | 0.47

2, | Cormocephalus 1.99 | 0.25 | 0.22 | 2.99 | 3.94 | 3.17 338] ]o.18}0.32]0.20 0.20 | 0.43
al paranetees

\o

154 Arkady A. Schileyko & Evgeniya N. Solovyeva/ ZooKeys 840: 133-155 (2019)

s[9 [o[n B16 [17

sic aa 1.76 | 0.50 | 0.65 | 2.92 | 3.92 | 3.15 0.26 | 0.19 | 0.42
monteithi
1.

01
Ee ee 30 | 2.80] 1.78 | 2.39 | 1. 02) 1. 0.23 | 0.31 | 0.41
octosulcatum

Ethmostigmus
rubripes

1.08 | 2.25 | 1.28] 2.04] 1. : : j . . . 0.25 | 0.39

Hemiscolopendra

: 1.70 | 0.84] 1.59 | 0. ; ‘ ; : , 1.91} 1.20 0.41
marginata

pial 3.31 | 2.37 | 3.10] 1. 93 | 3.01 | 3.49 | 2.52| 2.44 | 2.79 | 2.30 | 2.30
longitarsi
Newportia

A 3.28 | 2.09 | 2.65 } 1. ; f ; : , 2.70 | 2.04 | 2.09 | 0.56
monticola

piled aie 3.06 | 1.98 | 2.53 2.20] 1.76| 2.04 | 1.33
quadrimeropus

20 1.68 | 1.86 | 0.83 | 1.64 | 0.50 | 0.49 | 2.75 | 3.64 | 2.87 | 3.25 | 0.38 | 0.54] 1.74 | 1.25 | 0.60 | 2.44
21 1.41 | 2.10 | 1.08 | 1.65 | 3.13 | 3.98 | 3.14 | 3.52 | 1.55 | 1.54] 0.94 | 0.55 | 1.49 | 2.44

Appendix 6 (continuation)

Uncorrected p-distances (%) for sequences of 18S nuDNA gene for species (above diagonal). Standard

error estimates are shown above the diagonal.

8 [9 |B 31 | 2 [33

iT meee 0.29 | 0.34 | 0.31 0.36
chilensis

2 |Alipes grandidieri 0.33 | 0.37 | 0.32 0.35

elias ened 0.34 | 0.35 | 0.31 0.37
| formosus

Pea awe ce 0.27 | 0.15 0.31
brevicornis

5 |Asanada socotrana 0.33 | 0.27 0.35

6 Campylostigmus 0.31 | 0.24 0.39
decipiens

J | Carepioniayees 0.29 | 0.21 0.34
orientalis

8 | Cryptops australis 0.45 | 0.42 0.39

9 | Grypops spnipes 0.44 0.8

10: | Pers 0.44 | 0.45 | 0.42 | 0.41 0.38
trisulcatus
11 | Cryptops weberi 0.41 | 0.46 | 0.41 | 0.43 | 0.42 0.45
in| ee 0.24 | 0.30 | 0.21 | 0.24 | 0.37 0.34
aurantiipes
eae Hibesne aoa 0.28 | 0.19 | 0.23 | 0.38 0.31
monteithi
Plt ce uta 0.36 | 0.29 | 0.29 | 0.36 0.33
octosulcatum
0.31

ra mee eae 0.29 | 0.34 | 0.26 | 0.27 | 0.38
rubripes

tiles ea cate ao 0.22 | 0.26 | 0.19 | 0.21 | 0. 0.31
marginata

22 ;
Tales 0.19 | 0.32 | 0.43 | 0.39 | 0.40 | 0.40 | 0.40 | 0.40 0.39 | 0.34 | 0.39 | 0.38 | 0.36
longitarsi

18. | eee 0.34 | 0.37 | 0.31 | 0.36 | 0.35 33 | 0.30
monticola

foal eeeanee 0.35 | 0.36 | 0.33 | 0.36 | 0.34 31 | 0.29
quadrimeropus

Biel ene 2.09 | 2.04 0.26 | 0.25 | 0.24 | 0.25 | 0.23 | 0.28 | 0.18 | 0.21 | 0.37 | 0.32 | 0.26 | 0.29 | 0.29
glauerti

21 | Otostigmus astenus | 2.24 | 2.07 | 1.27 0.18 | 0.18 | 0.19 | 0.30 | 0.34 | 0.27 | 0.29 | 0.37 | 0.34 | 0.19 | 0.33 | 0.33

On the taxonomic position of the enigmatic genus Yonkinodentus Schileyko, 1992... 155
aD 2B Bs [sl [s
igs eee 2.04 | 1.93 | 1.14 055] [0.6] 0.17 0.29 0.38 | 0.33 | 0.15 | 0.33 | 0.31
caraibicus
Otostigmus
23 0.18 | 0.27 | 0.34 | 0.25 0.31
rugulosus
24 | Rhysida nuda 0.29 | 0.35 | 0.25 0.32
Dea lpre epee 0.27 | 0.17 0.30
morsitans
6 Scolopendra 0.23 0.35
cingulata
27 Scolopendra 1.10 0.28
subspinipes
28 | Scolopendra viridis| 2.31 | 2.10 | 0.76 | 1.60 | 1.36 | 1.41 | 1.36 | 1.20] 1.76] 0.87] | 0.40 | 0.32 | 0.27 | 0.30 | 0.31
29 | Scolepocryptops 2.48 | 3.01 | 2.37 | 2.76 0.34
mi1erstit
30, [ee epocopies 1.71 | 2.01 | 1.60 | 1.93 | 1.50 0.28
sexspinosus
ian tir tad 2.09 | 1.82 | 1.36 | 0.66 | 0.43 | 0.60 | 0.60 | 1.47 0.29
violaceus
go [ seers 2.18 | 1.60 | 1.93 | 2.34 1.77 0.20
erythrocephalus
33. | Theatops posticus 1.54 | 2.28 | 1.49 | 1.76 | 2.22 1.49 | 0.83