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\begin{document}

\title{New method for taxonomic descriptions with coded notation, producing
dynamic and interchangeable outputs}



\author[1]{Douglas Zeppelini}%
\author[1]{Misael Augusto de Oliveira Neto}%
\author[1]{João Victor Lemos Cavalcante de Oliveira}%
\author[1]{Aila Soares Ferreira}%
\author[1]{Roniere Andrade de Brito}%
\author[1]{Bruna Carolline Honório Lopes}%
\author[1]{Nathan Paiva Brito}%
\author[1]{Luis Carlos Stievano}%
\author[1]{Estevam Araujo de Lima}%
\affil[1]{Universidade Estadual da Paraiba}%


\vspace{-1em}



  
  \date{May 24, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
A new proposal for taxonomic species description is presented to replace
the traditional descriptive texts. This is an attempt to enhance the
species description rate and to make the description output available to
other scientific disciplines, machine learning, lucid identification
keys, big data analysis and its applications. The method consists in
presenting the description of the overall morphology in a coded matrix,
following a character list with detailed observed conditions for each
character. The method is supposed to be dynamic and open to amendments
and new data addition as they become available. We test the new method
describing five new species of Collembola Symphypleona of the genus
Pararrhopalites as a generalized model and made the coded output
available. We conclude that a coded taxonomic description is an advance
to the traditional taxonomic text, with potential to enhance the global
descriptions rate. The generated data is a dynamic matrix that can be
expanded with any data that becomes available, also it can be easily
used in other fields of science, allowing non-experts to access the data
for phylogenetic, biogeographic, ecological studies and big data
analysis. Furthermore, it is a step forward to a general template to
semi-automated taxon recognition and auxiliary tools for species
description using machine learning.%
\end{abstract}%



\sloppy


\textbf{Authors}

Douglas Zeppelini\selectlanguage{ngerman}¹\textsuperscript{,}²\textsuperscript{,*}, Misael
Augusto de Oliveira Neto¹\textsuperscript{,}², João Victor Lemos
Cavalcante de Oliveira¹, Aila Soares Ferreira¹, Roniere Andrade de
Brito¹, Bruna Carolline Honório Lopes¹\textsuperscript{,}², Nathan Paiva
Brito¹\textsuperscript{,}², Luis Carlos Stievano¹, Estevam Cipriano
Araujo de Lima¹.

¹Laboratório de Sistemática de Collembola e Conservação -- Instituto de
Biologia de solo -- Universidade Estadual da Paraíba, Campus V, João
Pessoa, PB 58070\selectlanguage{english}-450, Brazil.

2Programa de Pos\selectlanguage{english}-graduacao em Ciencias Biologicas -- Zoologia,
Universidade Federal da Paraiba, Joao Pessoa, PB, Brazil.

Correspondence

Douglas Zeppelini, Laboratorio de Sistematica de Collembola e
Conservacao -- Instituto de Biologia de solo -- Universidade Estadual da
Paraiba, Campus V, Joao Pessoa, PB 58070\selectlanguage{english}-450, Brasil

Email: zeppelini@daad-alumni.de

\textbf{Title}

\textbf{New method for taxonomic descriptions with coded notation,
producing dynamic and interchangeable outputs}

\textbf{Running Title}

CODED TAXONOMIC DESCRIPTION

\textbf{Abstract}

A new proposal for taxonomic species description is presented to replace
the traditional descriptive texts. This is an attempt to enhance the
species description rate and to make the description output available to
other scientific disciplines, machine learning, lucid identification
keys, big data analysis and its applications. The method consists in
presenting the description of the overall morphology in a coded matrix,
following a character list with detailed observed conditions for each
character. The method is supposed to be dynamic and open to amendments
and new data addition as they become available. We test the new method
describing five new species of Collembola Symphypleona of the
genus\emph{Pararrhopalites} as a generalized model and made the coded
output available. We conclude that a coded taxonomic description is an
advance to the traditional taxonomic text, with potential to enhance the
global descriptions rate. The generated data is a dynamic matrix that
can be expanded with any data that becomes available, also it can be
easily used in other fields of science, allowing non-experts to access
the data for phylogenetic, biogeographic, ecological studies and big
data analysis. Furthermore, it is a step forward to a general template
to semi-automated taxon recognition and auxiliary tools for species
description using machine learning.

\textbf{KEYWORDS}

Coded species description, Diversity decline, Taxonomy, Collembola,
Machine learning

\textbf{INTRODUCTION}

Taxonomy has been the focus of debate since the XIX century, and even
recently the recognition of the taxonomic research is subject of
discussion (Packer et al., 2018, Zeppelini et al., 2021). The global
diversity crisis exposes the urgency for investment in taxonomy to
reveal the largely unknown species diversity. Using Collembola as a
parameter, where about 20\% of its estimated diversity is known (Hopkin
1997), between 100 and 120 new species are described each year, and it
would take to taxonomists more than 400 years to uncover and describe
all the unknown species diversity (Potapov et al., 2020). To be able to
understand the diversification processes in Collembola, we need to speed
up the rates of species description. This is a matter of concern in
every area of entomology, and in some extent, the whole zoology.

Collembola Lubbock, 1870 are minute wingless arthropods, basal hexapods
found in every terrestrial habitat on the planet, including soil, leaf
litter, canopy trees and caves (Bellinger et al., 1996-2023, Hopkin
1997). There are about 9000 described species, and its diversity is
extensively underestimated and poorly known (Bellinger et al.,
1996-2023, Hopkin 1997). They play important role in the food web and
the global metabolism (Bardgett \& van der Putten 2014, Filser et al.,
2016, Potapov et al., 2023, Rusek 1998).

Similar to many other taxonomic groups of meso and micro fauna,
Collembola taxonomy is largely based on morphological analysis,
observing, and describing discrete variations in diagnostic characters.
The most abundant morphological source of information for species
definition in Collembola is the number, distribution, and shape of
cuticular chaetae, this is called chaetotaxy. The current morphological
approaches for inference of homology, chaetotaxic systems for chaetal
identification, are often room for great subjectivity depending on what
is seen and what is visible under an optic microscope, and often
different chaetotaxy systems are hardly comparable (Betsch 1997, Betsch
\& Waller 1994, Bretfeld 1990, 1999, Potapov et al., 2020). The
challenges and perspectives for Collembola taxonomy is discussed in
detail, and the need for an integrative taxonomy and international
efforts to direct financial support and expertise recognition to face
the global biodiversity crisis, was also the focus of debate (Potapov et
al., 2020, Zeppelini et al., 2021).

The impact of recent technologies of high-resolution imaging, molecular
sequencing and machine learning will be a great deal towards taxonomic
techniques that can improve new and known taxa recognition (Potapov et
al., 2020). Integrative taxonomy, combining morphological and molecular
data to define species limits is likely to be a trend for most taxonomic
groups, not only Collembola.

There is, however, a particular aspect in Collembola (and nearly every
taxon of the micro-fauna) that affects the viability of including
molecular sequences in new species descriptions, in many, if not most
cases. It is rather a logistic problem, but many times there is not an
alternative. The problem is that almost all new species are discovered
under light microscope, which means that the specimen was mounted in a
slide, after being cleared under several different techniques of
chemical washes, which destroy the tissues and, consequently, genetic
material.

It is only after the taxonomic identification, that a species is
recognized as new for science or undescribed. More often than not, the
material analyzed is a limited set of specimens, and there is no
available material for molecular analysis after the taxonomic
identification and morphologic study. Accepting that molecular analysis
facilities are available, many times the biological specimens needed for
molecular sequencing may be available only in a future, after the
species is described. Even when Scanning Electron Microscopy (SEM) is
possible, depending on the structure, it is hard to get images of all
diagnostic features and light microscopy may be needed as well. However,
high-resolution imaging and molecular data are powerful tools, and may
be indispensable for accurate taxonomic research and species
delimitation. Therefore, the morphologic descriptions must be dynamic,
open to easy amendment and additional data insertion. Furthermore, it
must be presented in an interchangeable language, to allow the
information to flow across different disciplines.

Among all methods applied to the external morphology study of
Collembola, chaetotaxy is certainly the most complex and extensively
detailed (Betsch \& Waller 1994, Cassagnau 1974, Deharveng 1983,
Fjellberg 1999, Jordana \& Baquero 2005, Nayrolles 1988, 1990a, 1990b,
Potapov 2001, Szeptycki 1979, 1972, Yosii 1960). There are many chaetae
and groups of chaetae that vary in position and shape in such a way that
they allow a great deal of homology inferences. However, the most
advanced approaches are also very complex, which makes interpretation
difficult and increases ambiguity. These aspects circumscribe the deep
taxonomic research to restricted groups of experts, posing difficulties
to comparative studies even among different Orders of Collembola. In
addition, the traditional descriptive texts with morphological and
chaetotaxic information are difficult to integrate with machine learning
and computational novelties, which could give a lot of agility to
phylogenetic analysis, big data comparison, biogeography, and their
various applications (Potapov et al. 2020).

Despite all advances in technological instruments and methods, taxonomic
descriptions are still written basically in the format as it was about
two centuries ago, with a hermetic language in nearly incomprehensible
texts for non-experts. This is often a greater barrier to communication
among different areas of science, than the access of high-tech equipment
and analytical facilities.

The proposal of a coded and illustrated description of new species that
can be easily imported, transformed, amended, corrected, or expanded is
presented as an alternative to the traditional descriptive taxonomic
method.

The strength of the coded description is that new characters, whether
morphological, molecular, ecological, can be easily added to the list
and improve the descriptive matrix as new information is produced. These
matrices can be uploaded to public libraries and kept up to date with
all available information about the species, and linked to data bases as
GBIF, ZooBank and electronic taxonomic catalogs available in different
parts of the world \emph{e.g., fauna.jbrj.gov.br/fauna/listaBrasil
(Zeppelini et al., 2023); www.collembola.org(Bellinger et al.,
1996-2023)} .

\textbf{MATERIALS AND METHODS}

\textbf{Coded Taxonomic Description}

The Order Symphypleona Borner, 1901 shows some ambiguity in current
morphological methods, particularly when describing the head and body
chaetotaxy (Betsch 1997, Betsch \& Waller 1994, Bretfeld 1990, 1999,
Christiansen \& Bellinger 1998). The Order is composed by springtails
with globular body shape, as result of modification and fusion of
thoracic and abdominal segments I-IV, this condition hinders the direct
assignment of segments identity.

An approach that can reduce the ambiguity of the taxonomic descriptions,
is the description of body parts into coded morphological units,
straightforwardly representing the actual body segments and appendicular
whorls (Hopkin 1997, Jura et al., 1987, Nayrolles 1988, 1990a, 1990b,
1991, Tomizuka \& Machida 2015), in such a way that any species can be
compared from the coded data base. This is in replacement to the
traditional descriptive text, many times with ambiguous terminology, and
often applying different and not directly comparable chaetotaxic
systems. The coded notation method would lead to a more comprehensive
analysis of the chaetotaxy, as well as a direct availability of the data
for comparative studies. Furthermore, a coded description can easily be
amended, molecular data can be added in the character list and matrix,
and new complementary morphological features can be inserted as new
information is available.

The qualitative description of the shape and size of the different
chaeta is also subject to a great deal of ambiguity and poor definition,
the adjectives are not standard and the very definition of what is a
macro, meso or microchaeta is not always clear. Therefore, a bank of
shapes with high quality images is imperative to discard all the
subjective descriptions. There are several chaetae banks published for
different groups of Collembola, including some with precise line
drawings(Betsch 1980, Christiansen 1966, Deharveng 1983, Nayrolles
1991), and some with SEM photography(Cipola et al., 2020, de Lima et
al., 2022, Luki\selectlanguage{polish}ć \selectlanguage{english}et al., 2010, Zeppelini et al., 2022, Zhang \&
Deharveng 2015), it is a matter of time to have a fully reliable chaetal
shapes collection, so a specific chaeta can be addressed directly by its
reference in the bank, in the coded description.

A standard, fully coded method for species description may be an
improvement to the traditional descriptive text, it may allow to use
machine learning and high-quality imaging to enhance the efficiency of
species descriptions and diversity recognition, offering a powerful tool
to understanding of global processes, and face the diversity decline.

\textbf{Chaetal fields and morphological units}

We attempt to access the chaetotaxy of the head and great abdomen of
Collembola, by identifying body segments arranged in each tagma (Figs.
1-2), or whorl in each appendage (Figs. 3-5) (Hopkin 1997, Jura et al.,
1987, Nayrolles 1988, 1990a, 1990b, 1991, Tomizuka \& Machida 2015).
Each segment has its own set of chaetae, and more than one chaetal field
may occur in a single body segment. A chaetal field is a group of
associated chaetae that are consistently observed in a given body
segment, often associated with some landmark on the cuticle (Figs. 6-7).

The morphological units are the actual observed character in a given
species. Once recognized all the chaetal fields, the characters are
listed, and its inherent character states are described. Each
morphological unit in the character list is given a code (0, 1, 2) to
each observed condition. It is important to note that it is not a
phylogenetic matrix, once the codes in the resulting matrix are not
supposed to include hypothesis of ordering or polarity, and both
apomorphies and plesiomorphies may be listed as character, instead, it
is a descriptive coded character state matrix.

\textbf{Head, body, and appendages chaetotaxy}

Chaetotaxic systems attempt to labeling each chaeta along the body,
where the label indicates a specific chaeta and its position in the body
of the animal, then a qualitative description is made (\emph{e.g} .
spinelike chaeta, macrochaeta, club shaped sensillum, palmate, serrate,
lanceolate), bringing subjectivity in the interpretation of a specific
chaeta labeling, and by the many different adjectives that can apply for
a given shape, depending on the author.

Here we map regions that can be compared in different taxonomic groups,
the chaetal fields, within each head and body segment (Fig. 1-2), and to
appendages (Figs. 3-5), and refer to a shape for the chaeta in an image
data set, the chaetae bank, with images of each kind of chaeta found in
the taxon (Fig. 8).

The chaetotaxy of individualized chaeta is replaced by a code describing
the total number of chaetae in the chaetal field, and the qualitative
description of the different kind of chaetae found in each chaetal
field, is replaced by the respective number in the chaetae bank that
represents the actual observed shape and size, to compose the
morphological unit definition (see Table 1).

\textbf{Coded descriptive dataset}

The information of the whole collection of data of each species, result
in a dataset as synthesized in table 1, this is the final morphologic
description outcome and represents the complete and up to date set of
information for each studied species. However, it is dynamic and open to
additional information, when available.

The coded dataset is hierarchically ordered in four columns namely,
Tagma, Segment, Chaetal field and Morphological unit (Table 1), a fifth
column is inserted with the coded information of the species (Fig. 1).
The lines of the resulting dataset bring the different features of the
chaetotaxy and general morphology (\emph{e.g.,} eyes, foot complex). The
cells in the column of morphological unit, are the actual features to be
observed in the specimen, where each one is a recognizable morphological
unit of the animal whole morphology, that is described and coded in the
character list (Appendix 1).

Using an example of a defined character and its code according to
appendix 1, we have: Antennal / ANT / ANT M + L / 3+1+3 4(9)3(7), were
Antennal indicates the segment, ANT is the chaetal field (see Fig. 1A),
ANT M+L is the morphologic unit, 3+1+3 are the observed chaetae in the
respective chaetal field (3 paired and 1 axial according to bilateral
simetry), and 4(9)3(10) means that there are four type ``9'' chaetae,
and three type ``7'' chaetae (total 7 chaetae or 3 +1+3), the types are
defined by comparison among the photos in figure 8. To the combination
of chaetae number and type is given a numeric code (this example is
coded 0 in the character list Appendix 1).

\textbf{Testing the coded description}

To test the proposed system, we describe five new species of the
genus\emph{Pararrhopalites} Bonet \& Tellez, 1947
(zoobank.org:pub:9ED865EA-F95A-4CBE-947C-3A5C6CD81907), from the Order
Symphypleona, using \emph{P. fallaciosus} \textbf{sp. n.} , as the
explanatory example in the SEM overall morphologic analysis. First, we
describe \emph{P. fallaciosus} \textbf{sp. n} ., where all the chaetal
fields are delimited, and the species is morphologically defined. The
character list is derived from this revision and presented in appendix
1. The final descriptions for the remaining new species will bring the
code only (Morphologic unit code).

Here we propose the coded description for \emph{Pararrhopalites} , a
genus of Symphypleona, however, once fully established, the system must
be applicable to the Orders Poduromorpha and Entomobryomorpha as well.
Ideally a similar approach could be used to any other zoological taxa.

To proceed the segmental delimitation and definition, we addressed the
results from developmental biology (Hopkin 1997, Jura et al., 1987,
Tomizuka \& Machida 2015), and the chaetal fields were defined using SEM
(Tescan Vegas III) to map the clusters of chaetae associated to
cuticular landmarks. Cephalic chaetotaxy, great abdomen chaetotaxy, and
small abdomen (abdominal segment VI) chaetotaxy followed the system
proposed by Betsch \& Waller (1994) and Betsch (1997). For the
appendages, we followed Nayrolles (1988, 1990a, 1990b, 1991) for
antennae, legs and furcula; and tenaculum after Richards (1968).

The chaetotaxic systems were used to verify the congruence of the
morphological landmarks and associated groups of chaetae which display
constant expression (\emph{i.e.,} chaetal fields). All chaetotaxic
information, the actual observed character condition, was described in
the character list, and coded accordingly (Appendix 1).

The descriptions are presented as a list of coded characters in the
descriptive plate of each newly described species. The exception is made
to \emph{P. fallaciosus} \textbf{sp. n.} , where the morphologic units
are described in the matrix corresponding to the chaetotaxy system cited
above, as an example of what the observed features are (before coding).
The detailed chaetotaxy analysis for all the five species described here
is available in the supplementary material.

\textbf{RESULTS}

\textbf{Coded Description of \emph{Pararrhopalites fallaciosus} sp. n.}

\textbf{Collembola Lubbock, 1870}

\selectlanguage{ngerman}\textbf{Symphypleona Börner, 1901}

\textbf{Sminthuroidea Bretfeld, 1994}

\textbf{Sminthuridae Lubbock, 1862}

\textbf{Sminthurinae Lubbock, 1862}

\textbf{Pararrhopalites Bonet \& Tellez, 1947 (=Neosphyrotheca Salmon,
1964)}

\emph{Pararrhopalites fallaciosus} \textbf{sp. n.}

zoobank.org:act:C9126B2C-EB0F-4AC3-BD10-066A54998541

(Figs. 1-8)

\textbf{Type material.} Holotype female in slide (\#20441/CRFS/UEPB):
Brazil, Minas Gerais state, Caeté municipality, F-29 SSH, 19°49'22.18''S
43°41'51.61''W, 26.IV.2022, Ativo ambiental \emph{et al.} coll.
Paratypes in slides (\#20439, \#20440/CRFS/UEPB): 2 females, same data
as holotype.

Additional materials https://doi.org/10.15468/pbxmgz

\textbf{Etymology.} \emph{Pararrhopalites fallaciosus} \textbf{sp. n.}is
a reference to the similarities with a \emph{P. queirozi} Brito, Lima \&
Zeppelini, 2019, that can lead to an elusive identification.

\textbf{Habitat and distribution}

The species was collected in drilling holes, occurring in the
Subterranean Shallow Habitat (SSH), its known distribution is restricted
to the type locality, despite the sampling efforts in nearby areas and
in the whole region, an important mining area which is being
consistently sampled in the last decade.

Good's Biogeographic zone 27(Culik \& Zeppelini 2003, Good 1974). The
climate according to Köppen's system is As (Köppen 1936, de Sá Júnior et
al., 2012, Shear 1966), presenting dry winters and wet summers, average
temperatures of 18°C during winter and 22°C in summer (valid for all the
five species described here).

\textbf{Remarks}

The new species resembles \emph{P. queirozi} in the shape of the subanal
appendage and cephalic chaetotaxy but can be clearly distinguished by
the number of subsegments of Ant. IV (eight in \emph{P. queirozi} , 10
in the new species), the presence of inner tooth in all ungues, and
apical filament exceeding the tip of the unguis in all three empodial
complexes in \emph{P. queirozi} . The new species is similar to \emph{P.
hermesi}\textbf{sp. n.} in the shape of subanal appendages, the lack of
inner tooth of all ungues. They differ by the number of eyes (1+1 in
\emph{P. fallaciosus} \textbf{sp. n.} and 0+0 in \emph{P. hermesi}
\textbf{sp. n.} ), number of subsegments in Ant. IV (10 and nine
respectively), the lack of corner tooth in all unguiculi and mucro with
inner lamella smooth in \emph{P. hermesi} \textbf{sp. n.} The reduced
number of chaetae on the dorsal posterior part of the great abdomen
(17+17) also differentiate \emph{P. fallaciosus} \textbf{sp. n.} from
other species of the queirozi-group (all the species which share the
same female subanal appendages).

\emph{Pararrhopalites hermesi} \textbf{sp. n.}

zoobank.org:act:554455DB-364D-4C48-A0A9-33CC578C4E02

(Figs. 8, 9-13)

\textbf{Type material.} Holotype female in slide (\#6040/CRFS/UEPB):
Brazil, Minas Gerais state, Nova Lima municipality, cave RM-33,
20°02'05.30''S, 43°59'40.20''W, 30.VI.2014, Bioespeleo \emph{et al.}
coll. Paratype in slide (\#6038/CRFS/UEPB): 1 female, Brazil, Minas
Gerais State, Nova Lima municipality, RM-33, 20°02'05.30''S,
43°59'40.20''W, 02.IV.2014, Bioespeleo \emph{et al.} coll. Paratype in
slide (\#6039/CRFS/UEPB): 1 female, same data as holotype. Paratype in
slide (\#6042/CRFS/UEPB): 1 male, Brazil, Minas Gerais State, Nova Lima
municipality, RM-03, 20°02'37.50''S 44°00'22.40''W, 31.III.2014,
Bioespeleo \emph{et al.} coll. Paratype in slide (\#18060/CRFS/UEPB): 1
female, Brazil, Minas Gerais State, Nova Lima municipality, RM-33,
20°02'05.30''S, 43°59'40.20''W, 21.X.2017, A. S. Reis \& R. Zampaulo
coll.

Additional materials\emph{https://doi.org/10.15468/pbxmgz}

\textbf{Etymology.} \emph{Pararrhopalites hermesi} \textbf{sp. n.} is
named after the Greek god Hermes, in allusion to the wings of his flying
shoes, quite similar in shape to the subanal appendages of this species.

\textbf{Habitat and distribution}

This species is known from caves and SSH in a range over 200 km across
different lithologies. It is a regionally widespread SSH species, but it
is not abundant, as there are less than 20 records of this species.

\textbf{Remarks}

This species is part of a group with a specific kind of subanal
appendages (number 28 in Fig. 8), which includes \emph{P. queirozi}
and\emph{P. fallaciosus} \textbf{sp. n.} The species has an intermediary
number of subsegments on Ant. IV (nine subsegments), has no eyes, lacks
the corner tooth in all unguiculi, and the inner lamella of the mucro is
smooth. This combination of features can easily differentiate the three
species.

Despite its wide distribution, \emph{P. hermesi} \textbf{sp. n.}presents
some features that may be indicative of its relation to the SSH
environment, for instance eye reduction, Ant. IV shorter (nine
subsegments), overall small body size, and the reduction of the corner
tooth and apical filament on unguiculus.

\emph{Pararrhopalites atypicus} \textbf{sp. n.}

zoobank.org:act:14BAAE02-F5FB-4169-BCE8-9AFC6DE4FBB1

(Figs. 8, 14-18)

\textbf{Type material.} Holotype female in slide (\#2142/CRFS/UEPB):
Brazil, Minas Gerais State, Mariana municipality, cave FN-05,
20°13'17.81''S 43°26'02.93''W, 12.XII.2012, Bioespeleo \emph{et
al.}coll.~ Paratypes in slides (\#2139, \#2140, \#2141, \#2143,
\#2144/CRFS/UEPB): 4 females and 1 male, same data as holotype.

Additional materials\emph{https://doi.org/10.15468/pbxmgz}

\textbf{Etymology.} \emph{P. atypicus} \textbf{sp. n.} is Latin for
unusual, as this is the only species of this region which lacks the
interantennal sensillar triangle, known so far.

\textbf{Habitat and distribution}

This species is restricted to a single area, there are only 10 records
for three small caves in the same iron rock formation. The species is
most likely distributed along the \emph{canga} , a SSH formation
resulting of weathering of the iron rock, that often connect caves in
the same lithology.

\textbf{Remarks}

This species resembles \emph{P. sideroicus} Zeppelini \& Brito, 2014
and\emph{P. ubiquum} Zeppelini, Lima \& Brito, 2018, in the shape of the
subanal appendages and general body chaetotaxy but differ from all the
species of the genus with records in this area by lacking the
interantennal sensillar triangle, this feature seems to be shared by
both the species of the queirozi-group and the ubiquum-group. The
presence of only three chaetae in the dorsal cephalic area DII is also
very unusual for the genus and the reduced number of chaetae in the
dorsal posterior part of the great abdomen can be diagnostic for this
species.

\emph{Pararrhopalites ritaleeae} \textbf{sp. n.}

zoobank.org:act:C531BF30-7489-449F-B0A1-2B4F1BBF67EF

(Figs. 8, 19-23)

\textbf{Type material.} Holotype female in slide (\#4942/CRFS/UEPB):
Brazil, Minas Gerais state, Itabirito municipality, cave VL-35,
20°18'28.30''S, 43°56'32.30''W, 18.VI.2013, Mascarenhas \emph{et
al.}coll. Paratypes in slides (\#2577, \#2587, \#2595/CRFS/UEPB): 3
females, Brazil, Minas Gerais state, Itabirito municipality, cave VL-35,
20°18'28.30''S, 43°56'32.30''W, 07-10.V.2013, Mascarenhas \emph{et
al.}coll. Paratypes in slides (\#4938, \#4940, \#4943/CRFS/UEPB): 2
female and 1 male, same data as holotype.

Additional materials\emph{https://doi.org/10.15468/pbxmgz}

\textbf{Etymology.} \emph{Pararrhopalites ritaleeae} \textbf{sp. n.} is
named \emph{in memoria} m after Rita Lee a Brazilian singer deceased
may, 2023.

\textbf{Habitat and distribution}

\emph{Pararrhopalites ritaleeae} \textbf{sp. n.} is the most abundant
species presented here, but its distribution is restricted to a group of
seven caves in the same iron rock formation in the type locality. It is
known only from caves, but surface samplings are needed to survey the
epigeic fauna.

\textbf{Remarks}

This species is also part of the ubiquum-group and resembles the other
species of the group by the shape of the subanal appendages and general
body chaetotaxy. The absence of eyes is observed in \emph{P. ubiquum} ,
and the apical filament of the unguiculus III exceeding the tip of the
unguis is similar to \emph{P. sideroicus} . The features that
differentiate \emph{P. ritaleeae} \textbf{sp. n.} are the mucro inner
lamella smooth, Ant IV with nine subsegments, unguiculus III apical
filament exceeding unguis tip, Abd. IV lateral chaetae lacking.

D. Dias Silva contributed with insights on the chaetal fields definition
and body chaetotaxy of \emph{P. fallaciosus} \textbf{sp. n.}

\emph{Pararrhopalites ironicus} \textbf{sp. n.}

zoobank.org:act:47BFB22F-967A-437D-A9FA-63F1675FB00B

(Figs. 8, 24-28)

\textbf{Type material.} Holotype female in slide (\#18545/CRFS/UEPB):
Brazil, Minas Gerais state, Barão dos Cocais municipality, cave
BRU-0028, 19°52'49.00''S, 43°26'13.50''W, 20-24.IX.2021, Spelayon
\emph{et al.} coll. Paratypes in slides (\#18555, \#18566/CRFS/UEPB): 2
females, same data as holotype. Paratypes in slides (\#19047, \#19048,
\#19059/CRFS/UEPB): 2 females and 1 male, Brazil, Minas Gerais state,
Barão dos Cocais municipality, cave MDIR-0028, 19°52'49.00''S,
43°26'13.50''W, 14-18.III.2022, Spelayon \emph{et al.} coll.

No additional materials available.

\textbf{Etymology.} \emph{Pararrhopalites ironicus} \textbf{sp. n.} ,
from iron and the Greek suffix ``\emph{icus} '' (belonging or related
to), in allusion to its relation to the iron rock.

\textbf{Habitat and distribution}

\emph{Pararrhopalites ironicus} \textbf{sp. n.} is a very restricted
species, represented by only six records from two caves in the same iron
rock formation, called \emph{Serra do Tamanduá} , the caves are 2700 m
away from each other, and connected by SSH. It is a rare species, likely
to be confined to the subterranean environment.

\textbf{Remarks}

This is one more species of the ubiquum-group, this species is very
similar to \emph{P. sideroicus} , \emph{P. ubiquum} , \emph{P.
atypicus}\textbf{sp. n.} , and \emph{P.} ritaleeae \textbf{sp. n.} in
the general body chaetotaxy. The main differences are to be found in the
dorsal cephalic chaetotaxy. \emph{Pararrhopalites ironicus} \textbf{sp.
n.} can be differentiated from those species by presenting only five
spines (type 8 in Fig. 8), the Ant. I with Only five chaetae, the apical
organ of Ant. III chaeta ape is a normal chaeta (type 21 in Fig. 8),
unguiculus III apical filament reaching but not exceeding the tip of the
unguis and lacking the corner tooth.

\textbf{DISCUSSION}

To access the global species diversity, it is mandatory to enhance the
description rates of new taxa, mainly where the diversity is least
known. A description protocol that can communicate the morphologic
characters in a coded notation, allows the application of new
technologies in the research and machine learning, which may be a major
turnover in the discipline, and affect the species description rates.
The scarcity of trained taxonomists and the hermeticity of the taxonomic
description manuscripts, are the biggest barriers to the advance of the
knowledge on the species diversity and evolutionary processes of
diversification, both clue elements to understand the global diversity
decline.

In the study of Collembola, as well as in many other groups, the
information content of a traditional taxonomic text is often difficult
to access and cannot be transported to analytical software without a
detailed revision of the species description, which many times demands
an expert in the taxon. The traditional format is also almost impossible
to be used for machine learning, as there are many differences in the
presentation of the data, that can make the comparison among different
manuscripts impossible to non-experts and to artificial intelligence.
The open character list of the coded description allows easy insertion
and correction of the information, and either the character lists, the
chaetae banks and the coded species descriptions are fully compatible
with technologies that work with data matrices.

Our results can be synthesized in the following conclusions:

1 -- The coded taxonomic description is a notation method that produces
interchangeable data, fully available for different scientific
disciplines. The data can be used by non-specialists for different
purposes in science.

2 -- The method makes it possible to add any source of new data to the
description when it became available. It is dynamic and open as a
continuous list of characters, the updating of the knowledge of a given
species is not dependent of a traditional taxonomic revision.

3 -- The method allows machine learning that can help to speed the
species description rates and taxon identification where they are least
known. This can be an important tool to fight global diversity crisis.

4 -- Coded description is meant to Collembola but may be applied to any
taxonomic group, reducing the ambiguity of narrative descriptions. When
it is widely used, the comparative analysis will be almost a
straightforward process.

\textbf{AUTHOR CONTRIBUTIONS}

\textbf{Douglas Zeppelini} : Conceptualization; investigation; writing -
original draft preparation; methodology; validation; visualization;
writing - review and editing; formal analysis; project administration;
data curation; supervision; resources. \textbf{Misael Augusto de
Oliveira Neto} : data curation, visualization. \textbf{João Victor Lemos
Cavalcante de Oliveira} : formal analysis; investigation, validation,
data curation. \textbf{Aila Soares Ferreira} : formal analysis;
investigation, validation, data curation. \textbf{Roniere Andrade de
Brito} : formal analysis; investigation, validation, data
curation.\textbf{Bruna Carolline Honório} \textbf{Lopes} : formal
analysis; investigation, validation, data curation. \textbf{Nathan Paiva
Brito} : investigation. \textbf{Luis Carlos Stievano} :
investigation.\textbf{Estevam Cipriano Araujo de Lima} : formal
analysis; investigation, validation, data curation.

\textbf{ACKNOWLEDGMENTS}

Senior author is granted by CNPq PQ\# 301460/2022-1. J.V.L.C. Oliveira
is granted Fapesq/PB\# 51686.872.33410.16082022. B.C.H. Lopes is granted
by CAPES\# 88882.440392/2019-01. M.A. Oliveira-Neto is granted by
CAPES\# 88887.821077/2023-00. N.P. Brito is granted by CAPES\#
88887.826124/2023-00. The study was partially funded by Brandt Meio
Ambiente Ltda. and Gerdau S.A. Diego Dias da Silva shared insights on
the chaetal fields definition.

\textbf{DATA AVAILABILITY STATEMENT}

The data that support the findings of this study are openly available in
Coleção de Referência de Fauna de Solo at
https://www.gbif.org/dataset/8cf92f87-eaee-4d1c-96a9-49c3f5998351, and
in the Support Information File.

\textbf{CONFLICT OF INTEREST STATEMENT}

The authors declare no conflict of interest.

\textbf{ORCID}

Douglas Zeppelini, https://orcid.org/0000-0002-9026-1129

Misael Augusto de Oliveira Neto, https://orcid.org/0000-0001-9249-9088

João Victor Lemos Cavalcante de Oliveira,
https://orcid.org/0000-0003-0433-5927

Aila Soares Ferreira, https://orcid.org/0000-0002-7544-3944

Roniere Andrade de Brito, https://orcid.org/0000-0002-4613-7673

Bruna Carolline Honório Lopes, https://orcid.org/0000-0002-8931-2443

Nathan Paiva Brito, https://orcid.org/0000-0001-9030-4339

Luis Carlos Stievano, https://orcid.org/0000-0001-7302-583X

Estevam Cipriano Araujo de Lima, https://orcid.org/0000-0002-1680-4818

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\textbf{TABLE 1} Morphologic description of \emph{P.
fallaciosus}\textbf{sp. n} . The table brings the state of each
morphologic unit observed in the species. The code for each state is
presented in the character list (Appendix 1). EN, eyes number; D, dorsal
head; C, clypeus; LR, labrum; Ant, antennal segment; MD, mandibular
segment; MX, maxilar segment; LB, labium; LBT, labial triangle; Th,
thorax; Abd, abdomen; SCX, subcoxa; CX coxa; TR, trochanter; FE, femur;
Tib, tibiotarsus; PT, pretarsus; TR, tenacular ramus; TC, tenacular
corpus; MN, manubrium; DN, dens; ov, oval organ; M, medial dorsal head;
L, lateral dorsal head; a, anterior; m, medial; md, mid-dorsal; p,
posterior; pd, postero-dorsal; pl, pre labral; v, ventral; d, dorsal; l,
lateral; ``-``, absent; ``+'', present, ``\textgreater{}'', bigger than;
``\textless{}'', shorter than. (N) Numbers in parentheses represent the
shape of the chaeta in the chaetae bank (Figure 8).\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
Tagma & Segment & Chaetal field & Morphologic unit & Morphologic unit
state\tabularnewline
& & & Pigments & -\tabularnewline
HEAD & Eye & Eyes number & EN & 1+1\tabularnewline
& Dorsal & D & DI M + L & 5+5 4(9)6(8)\tabularnewline
& & & DII M + L & 3+1+3 7(8)\tabularnewline
& & & DIII M + L & 0+0\tabularnewline
& Clypeal & C & CI & 3+3 6(10)\tabularnewline
& & & CII & 5+1+5 11(10)\tabularnewline
& & & CIII & 2+1+2 5(10)\tabularnewline
& & & CIV & 1+1+1 3(10)\tabularnewline
& Labral & LR & LR a + m + p + pl & 4+5+5+6 20(10)\tabularnewline
& Antennal & ANT & ANT M + L & 3+1+3 4(9)3(7)\tabularnewline
& Mandibular & MD & MDI (d+v) & 2+2+1+1ov 4(10)2(11)\tabularnewline
& & & MDII (d+v) & 8+8+2+2ov 16(10)4(11)\tabularnewline
& & & MDIII (d+v) & 10+10 20(10)\tabularnewline
& Maxilar & MX & DMX + MX & 2+2 4(10)\tabularnewline
& Labial & LB & LBT & 4+4 8(10)\tabularnewline
& & & LB & 2+2 4(10)\tabularnewline
BODY & Thorax & Thorax I & ThI & 0+0\tabularnewline
& & Thorax II & ThII & 1+1 2(24)\tabularnewline
& & Thorax III & ThIII & 3+3 2(24)2(26)2(23)\tabularnewline
& Abdomen & Abdomen I & AbdI a + m + p & 3+3 6(25)\tabularnewline
& & Abdomen II & AbdII a + m + p (d + l) & 8+8 8(25)8(26)\tabularnewline
& & Abdomen III & AbdIII a + m + p (d) & 10+10
8(25)6(26)6(31)\tabularnewline
& & & AbdIII a + m (l) & 5+5 10(26)\tabularnewline
& & Abdomen IV & AbdIV ad + md + pd & 17+17 4(25)30(27)\tabularnewline
& & & AbdIV (l) & 3+3 6(26)\tabularnewline
& & & BPI & 2+2 4(26)\tabularnewline
& & & BPII & 1+1 2(26)\tabularnewline
& & & BPIII l + m & 13+13 10(26)16(18)\tabularnewline
& & Abdomen V & AbdV a + m + p & 7+7 12(26)2(31)\tabularnewline
& & Abdomen VI & AbdVI as & 4+4 8(26)\tabularnewline
& & & AbdVI ms & 2+2 4(26)\tabularnewline
& & & AbdVI mps & 3+1+3 7(30)\tabularnewline
& & & AbdVI ps & 1+1+1 3(26)\tabularnewline
& & & AbdVI aai + ai & 8+8 16(26)\tabularnewline
& & & AbdVI mi & 3+3 6(26)\tabularnewline
& & & AbdVI mpi & 4+4 6(26)2(28)\tabularnewline
& & & AbdVI pi & 3+3 6(26)\tabularnewline
APPENDAGES & Antenna & Antenna I & AntI & 6 5(9)1(21)\tabularnewline
& & Antenna II & AntIIA & 7 6(9)1(21)\tabularnewline
& & & AntII -0+ & 3 3(9)\tabularnewline
& & & AntIIB & 4 4(9)\tabularnewline
& & Antenna III & AntIIIA & 8 2(6)1(2)1(21)1(20)3(9)\tabularnewline
& & & AntIII -0+ & 8 4(9)4(5)\tabularnewline
& & & AntIIIB & 4 4(9)\tabularnewline
& & Antenna IV & Subsegments & 10\tabularnewline
& & number of chaeta/whorl & AntIVAI + II + III & 34
1(3)1(1)2(4)6(10)24(5)\tabularnewline
& & & AntIVMA & 8 1(4)6(5)\tabularnewline
& & & AntIVM & 9 1(4)8(5)\tabularnewline
& & & AntIVMB & 8 1(4)7(5)\tabularnewline
& & & AntIVBA & 3 3(5)\tabularnewline
& & & AntIVBM & 3 3(5)\tabularnewline
& & & AntIVBB & 4 4(5)\tabularnewline
& Legs & Subcoxa & SCXI & 1 1(26)\tabularnewline
& & & SCXII & 1 1(26)\tabularnewline
& & & SCXIII & 2 2(26)\tabularnewline
& & Coxa & CXI & 1 1(26)\tabularnewline
& & & CXII & 3 3(26)\tabularnewline
& & & CXIII & 5 5(26)\tabularnewline
& & Trocanter & TRI & 5 5(26)\tabularnewline
& & & TRII & 6 6(26)\tabularnewline
& & & TRIII & 6 1(15)5(26)\tabularnewline
& & Femur & FEI & 15 15(26)\tabularnewline
& & & FEII & 14 14(26)\tabularnewline
& & & FEIII & 12 12(26)\tabularnewline
& & Tibiotarsus & TibI I & 9 1(13)8(14)\tabularnewline
& & & TibI II & 8 1(13)7(16)\tabularnewline
& & & TibI III & 8 1(13)7(16)\tabularnewline
& & & TibI IV & 14 1(13)13(16)?\tabularnewline
& & & TibI V & 8 1(13)7(16)\tabularnewline
& & & TibI.FS & 2 2(16)\tabularnewline
& & & TibI.FP & 3 3(16)\tabularnewline
& & & TibII I & 9 1(13)8(14)\tabularnewline
& & & TibII II & 8 1(13)7(16)\tabularnewline
& & & TibII III & 8 1(13)7(16)\tabularnewline
& & & TibII IV & 14 1(13)13(16)\tabularnewline
& & & TibII V & 8 1(13)7(16)\tabularnewline
& & & TibII FS & 3 3(16)\tabularnewline
& & & TibII FP & 3 3(16)\tabularnewline
& & & TibIII I & 9 1(13)8(14)\tabularnewline
& & & TibIII II & 8 1(13)7(16)\tabularnewline
& & & TibIII III & 8 1(13)7(16)\tabularnewline
& & & TibIII IV & 14 2(13)12(16)\tabularnewline
& & & TibIII V & 8 1(13)7(16)\tabularnewline
& & & TibIII FS & 3 3(16)\tabularnewline
& & & TibIII FP & 3 3(16)\tabularnewline
& & Pretarsus & PTI & 1+1 2(12)\tabularnewline
& & & PTII & 1+1 2(12)\tabularnewline
& & & PTIII & 1+1 2(12)\tabularnewline
& & Unguis & Unguis I inner tooth & -\tabularnewline
& & & Unguis I tunica & -\tabularnewline
& & & Unguis II inner tooth & -\tabularnewline
& & & Unguis II tunica & -\tabularnewline
& & & Unguis III inner tooth & -\tabularnewline
& & & Unguis III tunica & -\tabularnewline
& & Unguiculus & Unguiculus I filament & \textgreater{}\tabularnewline
& & & Unguiculus I tooth & +\tabularnewline
& & & Unguiculus II filament & \textgreater{}\tabularnewline
& & & Unguiculus II tooth & +\tabularnewline
& & & Unguiculus III filament & \textless{}\tabularnewline
& & & Unguiculus III tooth & +\tabularnewline
& & Collophore & Collophore & 1+1 2(19)\tabularnewline
& Tenaculo & Tenaculo & TR & 3+3 teeth\tabularnewline
& & & TC a + p & 2 2(18)\tabularnewline
& Furcula & Manubrio & MN & 5+5 10(16)\tabularnewline
& & Dente & DNI & 5 1(16)4(22)\tabularnewline
& & & DNII & 3 3(16)\tabularnewline
& & & DNIII & 3 3(16)\tabularnewline
& & & DNIV & 3 3(16)\tabularnewline
& & & DNV & 3 3(16)\tabularnewline
& & & DNVI & 2 2(16)\tabularnewline
& & & DNB & 4 4(16)\tabularnewline
& & & DNAnt & 4,2,2,1,1\tabularnewline
& & Mucro & Mucro & Both lamellae serrated\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\textbf{FIGURE 1} \emph{P. fallaciosus} \textbf{sp. n.} cephalic
chaetotaxy and descriptive table. A, dorsal cephalic schematic
chaetotaxy; B, ventral cephalic schematic chaetotaxy; C, dorsal and
ventral cephalic SEM. Abbreviations in the descriptive table as in Table
1.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{FIGURE 2} \emph{P. fallaciosus} \textbf{sp. n.} body chaetotaxy
and descriptive table. A, whole body schematic chaetotaxy; B,\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{FIGURE 3} \emph{P. fallaciosus} \textbf{sp. n.} antennal
chaetotaxy SEM and descriptive table. A, whole antennal chaetotaxy; B,
Ant. IV apical subsegment; C\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

\textbf{FIGURE 4} \emph{P. fallaciosus} \textbf{sp. n.} legs chaetotaxy
and descriptive table. A, leg I; B, leg II; C, leg III. For each
represented leg, the left side is a\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

\textbf{FIGURE 5} \emph{P. fallaciosus} \textbf{sp. n.} Abdominal
appendages chaetotaxy. A, ventral tube lateral view and anterior view;
B, tenaculum anterior view and lateral view; C, furcula schematic
chaetotaxy; D, furcula posterior view SEM; E, furcula anterior view SEM.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

\textbf{FIGURE 6} \emph{P. fallaciosus} \textbf{sp. n.} SEM of the head
showing the chaetotaxy and cuticular landmarks. A, dorsal view; B,
lateral view; C, labrum; D, labium and adjacent chaetae (oval organs in
blue).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image7/image7}

\end{center}
\end{figure}

\textbf{FIGURE 7} \emph{P. fallaciosus} \textbf{sp. n.} SEM of the body
showing general chaetotaxy and cuticular landmarks. A, thorax and abd.
I-II lateral view; B, whole abdomen dorsal view; C, whole abdomen
lateral view, D, abd. IV-VI lateral view; E, female anal valve posterior
view; F, female subanal appendages and genital opening.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image8/image8}

\end{center}
\end{figure}

\textbf{FIGURE 8} Chaetae bank, it is a collection of SEM photographs of
different chaetae found along the body, the chaetae are representatives
of their shapes and sizes, but not of their position in the body,
therefore a given shape and size represented by a photo in the bank may
occur in different parts of the body. A, habitus \emph{P.
fallaciosus}\textbf{sp. n.} ; B, chaetae bank. 1 -- Bulb sensillum
(usually found in the apex of ant IV); 2 -- Microsensillum (\emph{e.g.}
, Aai on the apical organ of Ant. III); 3 -- Macrochaeta, curved with
blunt tip (often associated to bulb sensillum on the apex of Ant. IV); 4
-- Bristlelike mesochaeta; 5 -- Smooth macrochaetae; 6 -- Microsensillum
with blunt apex (occurs in pairs on the Ant. III apical organ); 7 --
Semi globular microsensillum; 8 -- Spiniform mesochaeta 9 -- Smooth
curved mesochaeta; 10 -- Smooth stout macrochaeta; 11 -- Oval organ; 12
-- Pre tarsal spine (other groups present a microchaeta instead); 13 --
Slender spinelike mesochaeta; 14 -- Smooth stout mesochaeta; 15 --
Strongly spinelike macrochaeta (\emph{e.g.} , metatrochanteral spine);
16 -- Slender smooth macrochaeta; 17 -- Smooth strong macrochaeta
(similar to 10); 18 -- Slender smooth microchaeta; 19 -- Smooth stout
microchaeta; 20 -- Curved spinelike microchaeta; 21 -- Bristlelike
microchaeta; 22 -- Spinelike mesochaeta; 23 -- J-shaped blunt
mesosensillum; 24 -- Smooth mesochaeta; 25 -- Heavily spiniform
mesochaeta (similar to 8); 26 -- Slender smooth mesochaeta; 27 -- Smooth
broad macrochaeta (similar to 10 and 17, but longer); 28 -- Featherlike
macrochaeta (subanal appendage); 29 -- Broad lanceolate macrochaeta
(subanal appendages, sometimes ciliated); 30 -- Broad smooth
macrochaeta; 31 - Trichobothria\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image9/image9}

\end{center}
\end{figure}

\textbf{FIGURE 9} \emph{P. hermesi} \textbf{sp. n.} cephalic chaetotaxy
and descriptive table. A, dorsal cephalic schematic chaetotaxy; B,
ventral cephalic schematic chaetotaxy.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image10/image10}

\end{center}
\end{figure}

\textbf{FIGURE 10} \emph{P. hermesi} \textbf{sp. n.} body chaetotaxy and
descriptive table.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image11/image11}

\end{center}
\end{figure}

\textbf{FIGURE 11} \emph{P. hermesi} \textbf{sp. n.} antennal chaetotaxy
and descriptive table. A, whole antennal chaetotaxy; B, Ant. IV apical
subsegment; C, apical organ on Ant. III.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image12/image12}

\end{center}
\end{figure}

\textbf{FIGURE 12} \emph{P. hermesi} \textbf{sp. n.} legs chaetotaxy and
descriptive table. A, subcoxa, coxa, trochanter and femur, legs I-III
respectively; B, tibiotarsus and empodial complex, legs I-III
respectively.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image13/image13}

\end{center}
\end{figure}

\textbf{FIGURE 13} \emph{P. hermesi} \textbf{sp. n.} abdominal
appendages chaetotaxy. A, ventral tube lateral view; B, tenaculum
lateral view; C, furcula chaetotaxy\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image14/image14}

\end{center}
\end{figure}

\textbf{FIGURE 14} \emph{P. atypicus} \textbf{sp. n.} cephalic
chaetotaxy and descriptive table. A, dorsal cephalic schematic
chaetotaxy; B, ventral cephalic schematic chaetotaxy.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image15/image15}

\end{center}
\end{figure}

\textbf{FIGURE 15} \emph{P. atypicus} \textbf{sp. n.} body chaetotaxy
and descriptive table.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image16/image16}

\end{center}
\end{figure}

\textbf{FIGURE 16} \emph{P. atypicus} \textbf{sp. n.} antennal
chaetotaxy and descriptive table. A, whole antennal chaetotaxy; B, Ant.
IV apical subsegment; C, apical organ on Ant. III.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image17/image17}

\end{center}
\end{figure}

\textbf{FIGURE 17} \emph{P. atypicus} \textbf{sp. n.} legs chaetotaxy
and descriptive table. A, subcoxa, coxa, trochanter and femur, legs
I-III respectively; B, tibiotarsus and empodial complex, legs I-III
respectively.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image18/image18}

\end{center}
\end{figure}

\textbf{FIGURE 18} \emph{P. atypicus} \textbf{sp. n.} abdominal
appendages chaetotaxy. A, ventral tube lateral view; B, tenaculum
lateral view; C, furcula chaetotaxy and mucronal lamellae.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image19/image19}

\end{center}
\end{figure}

\textbf{FIGURE 19} \emph{P. ritaleeae} \textbf{sp. n.} cephalic
chaetotaxy and descriptive table. A, dorsal cephalic schematic
chaetotaxy; B, ventral cephalic schematic chaetotaxy.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image20/image20}

\end{center}
\end{figure}

\textbf{FIGURE 20} \emph{P. ritaleeae} \textbf{sp. n.} body chaetotaxy
and descriptive table.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image21/image21}

\end{center}
\end{figure}

\textbf{FIGURE 21} \emph{P. ritaleeae} \textbf{sp. n.} antennal
chaetotaxy and descriptive table. A, whole antennal chaetotaxy; B, Ant.
IV apical subsegment; C, apical organ on Ant. III.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image22/image22}

\end{center}
\end{figure}

\textbf{FIGURE 22} \emph{P. ritaleeae} \textbf{sp. n.} legs chaetotaxy
and\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image23/image23}

\end{center}
\end{figure}

\textbf{FIGURE 23} \emph{P. ritaleeae} \textbf{sp. n.} abdominal
appendages chaetotaxy. A, ventral tube lateral view; B, tenaculum
lateral view; C, furcula chaetotaxy and mucronal lamellae. Circle with
an underline means chaeta present or absent.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image24/image24}

\end{center}
\end{figure}

\textbf{FIGURE 24} \emph{P. ironicus} \textbf{sp. n.} cephalic\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image25/image25}

\end{center}
\end{figure}

\textbf{FIGURE 25} \emph{P. ironicus} \textbf{sp. n.} body chaetotaxy
and descriptive table.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image26/image26}

\end{center}
\end{figure}

\textbf{FIGURE 26} \emph{P. ironicus} \textbf{sp. n.} antennal
chaetotaxy and descriptive table. A, whole antennal chaetotaxy; B, Ant.
IV apical subsegment; C, apical organ on Ant. III.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image27/image27}

\end{center}
\end{figure}

\textbf{FIGURE 27} \emph{P. ironicus} \textbf{sp. n.} legs chaetotaxy
and descriptive table. A, subcoxa, coxa, trochanter and femur, legs
I-III respectively; B, tibiotarsus and empodial\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image28/image28}

\end{center}
\end{figure}

\textbf{FIGURE 28} \emph{P. ironicus} \textbf{sp. n.} abdominal
appendages chaetotaxy. A, ventral tube lateral view; B, tenaculum
lateral view; C, furcula chaetotaxy and mucronal lamellae. Circle with
an underline means chaeta present or absent.

\textbf{APPENDIX 1}

The following list of characters is presented in a sequence from the
first cephalic segment to the sixth abdominal segment. It is a perfect
match of Table 1 and is also represented by an individualized table in
each species description plate. This list presents the observed
conditions of each character included in our study, a total of 115
morphological characters are listed.

\textbf{Color pattern}

Pigments:

0 -- Absent

1 -- Present

Body color:

0 -- White

1 -- Pale yellow

\textbf{Head}

Eye number:

0 -- 1+1

1 -- 0+0

2 -- 2+2

\textbf{Dorsal cephalic chaetotaxy}

Area DI total chaetae M + L:

0 -- 5+5 4(9)6(8)

1 -- Different combination

Area DII total chaetae M + L:

0 -- 3+1+3 7(8)

1 -- 2+1+2 5(8)

2 -- 1+1+1 3(8)

Area DIII total chaetae M + L:

0 -- 0+0

1 -- Different combination

\textbf{Clypeal chaetotaxy}

Clypeal area CI:

0 -- 3+3 6(10)

1 -- Different combination

Clypeal area CII:

0 -- 5+1+5 11(10)

1 -- Different combination

Clypeal area CIII:

0 -- 2+1+2 5(10)

1 -- Different combination

Clypeal area CIV:

0 -- 1+1+1 3(10)

1 - Different combination

\textbf{Labral chaetotaxy}

Labral formula a + m + p + pl:

0 -- 4+5+5+6 20(10)

1 -- Different combination

\textbf{Antennal area}

Antennal area chaetotaxy M + L:

0 -- 3+1+3 4(9)3(7)

1 -- 2+2 4(9)

\textbf{Mandibular segment chaetotaxy}

Area MDI (dorsal+ventral):

0 -- 2+2+1+1ov 4(10)2(11)

1 -- Different combination

Area MDII (d+v):

0 -- 8+8+2+2ov 16(10)4(11)

1 -- 12+12 2+2ov 24(10)4(11)

Area MDIII (d+v):

0 -- 10+10 20(10)

1 -- 9+9 18(10)

\textbf{Maxilar segment chaetotaxy}

Areas DMX and MX total chaetae

0 -- 2+2 4(10)

1 -- 1+1 2(10)

\textbf{Labial segment chaetotaxy}

Labial triangle total chaetae:

0 -- 4+4 8(10)

1 -- Different combination

Labial remaining chaetae:

0 -- 2+2 4(10)

1 -- Different combination

\textbf{Body}

\textbf{Thoracic chaetotaxy}

Thorax I total chaetae:

0 -- 0+0

1 -- at least 1+1

Thorax II total chaetae:

0 -- 1+1 2(24)

1 -- 0+0

2 -- Different combination

Thorax III total chaetae:

0 -- 3+3 2(24)2(26)2(23)

1 -- Different combination

\textbf{Abdominal chaetotaxy}

Abdome I a + m + p total chaetal:

0 -- 3+3 6(25)

1 -- 3+3 4(25)2(26)

Abdome II a + m + p (d + l) total chaetae:

0 -- 8+8 8(25)8(26)

1 -- 7+7 8(25)6(26)

Abdome III a + m + p (dorsal) total chaetae:

0 -- 10+10 8(25)6(26)6(31)

1 -- Different combination

Abdomen III a + m (lateral) total chaetae:

0 -- 5+5 10(26)

1 -- 5+5 8(26)2(23)

Abdomen IV ad + md + pd total chaetae (dorsal):

0 -- 17+17 4(25)30(27)

1 -- 27+27 4(25)50(27)

2 -- 28+28 56(27)

3 -- 30+30 60(27)

Abdomen IV total chaetae (lateral):

0 -- 3+3 6(26)

1 -- 2+2 4(26)

2 -- 0+0

Abdomen IV area BPI total chaetae:

0 -- 2+2 4(26)

1 -- Different combination

Abdomen IV area BPII total chaetae:

0 -- 2+2 4(26)

1 -- 1+1 2(26)

2 -- 0+0

Abdomen area BPIII l + m total chaetae:

0 -- 13+13 10(26)16(18)

1 -- 13+13 6(26)20(18)

2 -- 13+13 6(17)16(26)4(18)

Abdomen V a + m + p total chaetae:

0 -- 7+7 12(26)2(31)

1 -- 6+6 10(26)2(31)

Abdomen VI \textbf{as} total chaetae:

0 -- 3+3 6(26)

1 -- 4+4 8(26)

2 -- 6+6 12(26)

3 -- 7+7 14 (26)

Abdomen VI \textbf{ms} total chaetae:

0 -- 2+2 4(26)

1 -- Different combination

Abdomen VI \textbf{mps} total chaetae:

0 -- 3+1+3 7(30)

1 -- Different combination

Abdomen VI \textbf{ps} total chaetae:

0 -- 1+1+1 3(26)

1 -- Different combination

Abdomen VI \textbf{aai} + \textbf{ai} total chaetae:

0 -- 8+8 16(26)

1 -- Different combination

Abdomen VI \textbf{mi} total chaetae:

0 -- 3+3 6(26)

1 -- Different combination

Abdomen VI \textbf{mpi} total chaetae:

0 -- 4+4 6(26)2(28)

1 -- 5+5 8(26)2(28)

2 -- 4+4 6(26)2(29)

3 -- 5+5 8(26)2(29)

Abdomen VI \textbf{pi} total chaetae:

0 -- 3+3 6(26)

1 -- Different combination

\textbf{Appendages}

\textbf{Antenna}

Antenna I total chaetae:

0 -- 6 5(9)1(21)

1 -- 5 4(9)1(21)

Antenna II whorl \textbf{A} total chaetae:

0 -- 7 6(9)1(21)

1 -- 7 2(20)1(21)4(9)

2 -- 8 2(20)1(21)5(9)

Antenna II whorls \textbf{-} , \textbf{0} and \textbf{+} total chaetae:

0 -- 3 3(9)

1 -- 3 1(20)2(9)

2 -- 4 1(20)3(9)

Antenna II whorl \textbf{B} total chaetae:

0 -- 4 4(9)

1 -- Different combination

Antenna III whorl \textbf{A} total chaetae:

0 -- 8 2(6)1(2)1(21)1(20)3(9)

1 -- 8 2(6)1(2)1(21)2(20)2(9)

2 -- 8 2(6)1(2)2(20)3(9)

3 -- 8 2(6)1(2)5(9)

Antenna III whorls \textbf{-} , \textbf{0} and \textbf{+} total chaetae:

0 -- 8 4(9)4(5)

1 -- 9 1(9)8(10)

2 -- 9 9(10)

3 -- 10 10(10)

Antenna III whorl \textbf{B} total chaetae:

0 -- 4 4(9)

1 -- 5 5(9)

Antenna IV subsegments:

0 -- 10 subsegments

1 -- 9 subsegments

2 -- Less than 9 subsegments

3 -- More than 10 subsegments

Antenna IV \textbf{AI} + \textbf{AII} + \textbf{AIII} total chaetae:

0 -- 34 1(3)1(1)2(4)6(10)24(5)

1 -- 32 1(3)1(1)2(4)8(10)20(5)

2 -- 31 1(3)1(1)5(10)24(5)

3 -- 30 1(3)1(1)8(10)20(5)

Antenna IV \textbf{MA} total chaetae per whorl:

0 -- 8 1(4)7(5)

1 -- 9 1(4)8(5)

2 -- 10 2(4)8(5)

Antenna IV \textbf{M} total chaetae per whorl:

0 -- 9 1(4)8(5)

1 -- 10 2(4)8(5)

Antenna IV \textbf{MB} total chaetae per whorl:

0 -- 8 1(4)7(5)

1 -- 9 1(4)8(5)

Antenna IV \textbf{BA} total chaetae:

0 -- 3 3(5)

1 -- 5 5(5)

2 -- 6 6(5)

Antenna IV \textbf{BM} total chaetae:

0 -- 3 3(5)

1 -- 4 4(5)

2 -- 5 5(5)

3 -- 6 5(5)1(4)

Antenna IV \textbf{BB} total chaetae:

0 -- 4 4(5)

1 -- Different combination

\textbf{Legs}

Subcoxa I total chaetae:

0 -- 1 1(26)

1 -- Different combination

Subcoxa II total chaetae:

0 -- 1 1(26)

1 -- Different combination

Subcoxa III total chaetae:

0 -- 2 2(26)

1 -- Different combination

Coxa I total chaetae:

0 -- 1 1(26)

1 -- Different combination

Coxa II total chaetae:

0 -- 3 3(26)

1 -- Different combination

Coxa III total chaetae:

0 -- 5 5(26)

1 -- Different combination

Trochanter I total chaetae:

0 -- 5 5(26)

1 -- Different combination

Trochanter II total chaetae:

0 -- 6 6(26)

1 -- 5 5(26)

Trochanter III total chaetae:

0 -- 6 1(15)5(26)

1 -- Different combination

Femur I total chaetae:

0 -- 15 15(26)

1 -- 14 14(26)

2 -- 12 12(26)

3 -- 10 10(26)

Femur II total chaetae:

0 -- 15 13(26)2(13)

1 -- 14 14(26)

2 -- 12 12(26)

Femur III total chaetae:

0 -- 18 18(26)

1 -- 14 13(26)1(21)

2 -- 14 14(26)

3 -- 12 12(26)

Tibiotarsus I whorl I total chaetae:

0 -- 9 1(13)8(14)

1 -- Different combination

Tibiotarsus I whorl II total chaetae:

0 -- 8 1(13)7(16)

1 -- 9 1(13)8(16)

Tibiotarsus I whorl III total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus I whorl IV total chaeta:

0 -- 14 1(13)13(16)

1 -- 12 1(13)11(16)

Tibiotarsus I whorl V total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus I \textbf{FS} area total chaetae:

0 -- 2 2(16)

1 -- Different combination

Tibiotarsus I \textbf{FP} area total chaetae:

0 -- 3 3(16)

1 -- Different combination

Tibiotarsus II whorl I total chaetae:

0 -- 9 1(13)8(14)

1 -- Different combination

Tibiotarsus II whorl II total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus II whorl III total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus II whorl IV total chaetae:

0 -- 14 1(13)13(16)

1 -- Different combination

Tibiotarsus II whorl V total chaeta:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus II \textbf{FS} area total chaetae:

0 -- 3 3(16)

1 -- Different combination

Tibiotarsus II \textbf{FP} area total chaetae:

0 -- 3 3(16)

1 -- Different combination

Tibiotarsus III whorl I total chaetae:

0 -- 9 1(13)8(14)

1 -- Different combination

Tibiotarsus III whorl II total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus III whorl III total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus III whorl IV total chaetae:

0 -- 14 2(13)12(16)

1 -- Different combination

Tibiotarsus III whorl V total chaetae:

0 -- 8 1(13)7(16)

1 -- Different combination

Tibiotarsus III \textbf{FS} area total chaetae:

0 -- 3 3(16)

1 -- Different combination

Tibiotarsus III \textbf{FP} area total chaetae:

0 -- 3 3(16)

1 -- Different combination

Pretarsus I pre tarsal chaetae:

0 -- 1+1 2(12)

1 -- Different combination

Pretarsus II pre tarsal chaetae:

0 -- 1+1 2(12)

1 -- Different combination

Pretarsus III pre tarsal chaetae:

0 -- 1+1 2(12)

1 -- Different combination

Unguis I inner tooth:

0 -- present

1 -- absent

Unguis I tunica:

0 -- absent

1 -- present

Unguis II inner tooth:

0 -- present

1 -- absent

Unguis II tunica:

0 -- absent

1 -- present

Unguis III inner tooth:

0 -- present

1 -- absent

Unguis III tunica:

0 -- absent

1 -- present

Unguiculus I apical filament:

0 -- exceeding the unguis tip

1 -- reaching the unguis tip

Unguiculus I empodial tooth:

0 -- present

1 -- absent

Unguiculus II apical filament:

0 -- exceeding the unguis tip

1 -- reaching the unguis tip

Unguiculus II empodial tooth:

0 -- present

1 -- absent

Unguiculus III apical filament:

0 -- exceeding the unguis tip

1 -- reaching the unguis tip

2 -- shorter than unguis tip

Unguiculus III empodial tooth:

0 -- present

1 -- absent

Collophore total chaetae:

0 -- 1+1 2(19)

1 -- different combination

Tenaculum rami:

0 -- 3+3 teeth

1 -- different combination

Tenaculum Corpus (a+p):

0 -- 2 2(18)

1 -- different combination

Manubrium total chaetae:

0 -- 6+6 12(16)

1 -- 5+5 10(16)

2 -- 4+4 8(16)

3 -- 3+3 4(16)2(22)

Dental whorl I total chaetae:

0 -- 5 1(16)4(22)

1 -- different combination

Dental whorl II total chaetae:

0 -- 3 3(16)

1 -- 3 1(16)2(22)

Dental whorl III total chaetae:

0 -- 3 3(16)

1 -- 3 1(16)2(22)

Dental whorl IV total chaetae:

0 -- 3 3(16)

1 -- 2 2(22)

Dental whorl V total chaetae:

0 -- 3 3(16)

1 -- 2 1(16)1(22)

Dental whorl VI total chaetae:

0 -- 2 2(16)

1 -- different combination

Dental basal area \textbf{B} total chaetae:

0 -- 4 4(16)

1 -- different combination

Dental anterior chaetotaxy:

0 -- 4,2,2,1,1

1 -- different combination

Mucro lamellae:

0 -- Both lamellae serrated

1 -- Internal lamella smooth

\selectlanguage{english}
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\end{document}

