ABSTRACT A hotspot of subterranean Collembola in ferruginous rock caves and Mesovoid Shallow Substratum is revealed by the analysis of pseudocryptic diversity. The diversity is accessed by

detailed description of chaetotaxy and slight variation in morphology of 11 new species of _Trogolaphysa_ Mills, 1938 (Collembola, Paronellidae, Paronellinae) and the 50 previously recorded

species of springtails from caves, using optical and electronic microscopy. When combined with recent subterranean surveys, our results show an important reservoir of cave diversity in the

Mesovoid Shallow Substratum. Contrastingly the conservation policy for subterranean fauna in metallogenic areas in Brazil prioritizes the caves instead the cave species, which may be

POLAR ECOSYSTEMS Article Open access 09 February 2024 SUBSISTENCE OF EARLY ANATOMICALLY MODERN HUMANS IN EUROPE AS EVIDENCED IN THE PROTOAURIGNACIAN OCCUPATIONS OF FUMANE CAVE, ITALY Article

Open access 07 March 2023 INTRODUCTION Some areas are subject of intense fauna diversification, the term “hotspot” is used to indicate relatively small areas with high and exclusive

diversity, though there are different interpretations about what is the threshold which defines such an area. Hotspots may be defined by combining the richness, endemism, extension, and

threats to the area in focus1, however some approaches to subterranean fauna use an arbitrary cutoff of 20 restricted endemic species with no regards to environmental threats2. More recently

two South American subterranean hotspots were defined based in the richness of restricted endemic fauna, and fully addressed the conservation aspects of the surroundings of the cave

systems3. The species diversity of cave restricted fauna, with limited subterranean distribution and some degree of troglomorphism, known as troglobites4, is positively correlated to the

extension of the cave and the presence of perennial pools, and sometimes negatively correlated to the presence of streams, which can cause disturbance in the habitats and import a more

diverse troglophile fauna5. Therefore, the more diverse troglobitic fauna is supposed to be found in larger caves, which are often formed in limestone rock, this is corroborated by the cave

fauna hotspots found in limestone cave systems in Southeastern and Northeastern Brazil3,6. However, there are different shallow subterranean habitats (SSH), which are spaces that extend

through and across the soil and weathered rock matrix7. The mesovoid shallow substratum (MSS)7,8,9 seems to be the SSH that best fits the characteristics observed in ferruginous subterranean

habitats, even though it differs from the exact original definition for MSS8. These underground spaces connect, and somehow extends the cave habitat far beyond the human reach and might as

well be a climate refuge to epigeic fauna10,11. The Brazilian States of Minas Gerais (Southeast) and Para (North) represent the more important metallogenic areas in the country and

concentrate the mining activities and commodities production. The iron ore lithology presents a profusion of small and shallow caves, subterranean spaces and crevices that functions as

MSS12, providing habitat for a variety of species, including troglobites13,14. Previous studies found higher average relative richness, and distinctiveness in ferruginous rock12, than in

other lithologies. Unpublished data from caves and MSS in ferruginous rock, brings 87 morphotypes of Collembola with some degree of troglomorphism, most of them potential new species, from

Minas Gerais (73) and Para (14). A total of 38 species have been described so far (32 from Minas Gerai_s_, six from Para), including 22 troglobites (Table 1). Recent studies in Brazil have

surveyed hundreds of cave species, from sponges to vertebrates, more than 250 already described3,14,15,16,17, great part of it focused on large caves in karstic lithology6,18,19. Here we

present a group of 50 known species of Collembola found in Brazilian caves in different lithologies and add 11 new species of the genus _Trogolaphysa_ Mills, 1938 with some degree of

troglomorphism, from caves and MSS in ferruginous and limestone rock. The genus _Trogolaphysa_ has 69 described species worldwide, only eight have been recorded from Brazil so far: _T.

aelleni_ Yoshii, 1988; _T. ernesti_ Cipola & Bellini, 2017; _T. formosensis_ Silva & Bellini, 2015; _T. hauseri_ Yoshii, 1988; _T. hirtipes_ (Handschin, 1924), _T. millsi_ Arlé,

1939; _T. piracurucaensis_ Nunes & Bellini, 2018; and _T. tijucana_ (Arlé & Guimarães, 1979). Our results depict an important hotspot for cave Collembola in the State of Minas

Gerais, also corroborate the expected high species richness in ferruginous rock caves and MSS and shed some light to the impact of the MSS in the conservation policy as a refuge for

subterranean diversity. RESULTS This study presents 11 new species of cave _Trogolaphysa_ (Table 1), two new species from Para, eight from Minas Gerais from caves in metallogenic rock, and

one new species from Sao Paulo, found in a limestone cave. Species were collected directly from organic debris in caves, the MSS was accessed through samplings in drilling holes. These

results represent an increase of 25% in the previous 32 species of cave Collembola described for the State of Minas Gerais. Data from Para are still scarce, with only five previous records

from iron rock caves and one from sandstone cave (Table 1). The species from Sao Paulo is from a different lithology, a much larger cave with narrower connections to the MSS. It represents a

new record to add to the 17 known cave springtails from limestone caves in Brazil (Table 1). There are 50 valid species of cave Collembola previously recorded from Brazil, 38 troglobites

and 12 troglophiles. For limestone caves, there are 17 species from five different States at South and Southeastern Regions, all troglobites. There is a troglobitic species from sandstone

cave in the State of Para at Northern Region, and one troglobite from granitic caves in Minas Gerais. All the 34 remaining records are from ferruginous rock caves and MSS, 29 from the

Southeastern State of Minas Gerais (19 troglobites and 10 troglophiles), and five from Para (three troglobites and two troglophiles). With the results presented here the total number of cave

Collembola recorded for Brazil rises to 61 species, with 18 species from limestone caves, one species from sandstone cave, one from granitic cave, and iron caves and MSS with 44 known

subterranean species. The State of Minas Gerais present the highest richness for cave Collembola in Brazil, with five species from limestone and one from granitic caves, and 37 records of

species endemic to ferruginous rock shallow caves and MSS in Minas Gerais, this has important implications for the conservation areas policy in Brazil, which may apply to other ferruginous

rock subterranean environments in tropical areas in the world. DISCUSSION FERRUGINOUS MESOVOID SHALLOW SUBSTRATUM The iron ore deposits in Brazil present a semi continuous covering layer of

fragmented hematite and lesser components cemented by limonite, called _Canga_. It is formed by weathering and lixiviation, and produce a labyrinthic complex of subterranean spaces,

crevices, and tiny underground connections, depicting a habitat that is analogous to the MSS12. In temperate zones the MSS plays a role as refuge for arthropod fauna, mainly at high

altitudes where the cold weather can eliminate all the ectothermic fauna from the surface10,11. Similarly, seasonal migration movements are observed in the MSS for different taxa as response

to hot dry summer20,21. When troglobitic fauna is concerned the MSS has a different role, cave restricted Collembola showed higher underground dispersal capacity than troglophiles22,

therefore, the MSS can connect neighboring caves systems and extend their distribution range. In Brazilian metallogenic rock, cave species richness is higher than in any other lithology12.

The cave Collembola found in Brazil corroborates this assumption, from the total of 61 known cave species, one troglobite was recorded from sandstone caves and one from granitic cave, 18

species were recorded from limestone caves (all troglobites), and 44 species from iron caves and MSS (31 troglobites). Three troglobitic species were recorded from both limestone and iron

caves, in both cases the caves are separated by large distances and the lithologies are disjunct. This incongruent and disjunct distribution is an indication of potentially unrecognized

cryptic or overlooked species. This is more relevant when considered that ferruginous rock represents only 0.15% of the Brazilian territory (carbonatic rock 3.1%), nearly 10,000 km2

(carbonatic rock 260.800 km2, Brazilian territory 8.516.000km2)23, and that most of the biospeleological research is focused on large caves, usually in limestone6,18,19. The high richness of

species restricted to small shallow caves, indicates that MSS plays a role as an extension of the cave environment. The State of Minas Gerais is the most diverse with 40 species of cave

Collembola, the complex mosaicist lithology and the ecotone Cerrado Forest-Atlantic Forest are the main barriers associated to the richness of species restricted to caves and MSS. In this

State, the iron rock subterranean habitats host 29 troglobites and provide habitat and refuge to 11 known troglophiles species. For caves in non-ferruginous lithologies, the size and number

of entrances influence the species richness by giving the surface fauna access to the subterranean environment, and as a sink for organic matter input5. Contrastingly the caves in iron rock

are small and shallow, often with few meters of horizontal development, the connections to the MSS are conspicuous and abundant, providing a rather continuous subterranean habitat. In this

context, instead, the distribution of the troglobitic species suggests that the entrances of iron rock caves are the limits of the available subterranean habitat for troglobites inside-out,

and of suitable habitats for troglophiles outside-in (Fig. 1). We can consider the entrances of these caves as windows of the MSS, the spatial limit of the subterranean environment which

presents the minimum conditions to the survival of a troglobite, while partially inhibits the dispersion of troglophiles deeper in the MSS. Troglobites can disperse underground more

efficiently than troglophiles, however troglophiles are more efficient than troglobites to disperse through the surface22. In ferruginous lithology the size of the cave and its entrance

influences the species richness5, mainly because large iron caves can greatly affect the capacity of collectors and biologists to access the troglobitic fauna in the MSS, as the number of

accessible connections to the MSS increases exponentially with the length of the cave in iron rock12. Another contrast of ferruginous rock caves is that the biotrophic flow seems to be

inverted (Fig. 1), in limestone caves the energy and fauna come from outside mostly through the cave entrance, and the fauna eventually speciate to become troglobitic, possibly restricted to

the depths of a single cave. Despite the demonstrated existence of an epikarst, the particularities of the weathering process, the water percolation24, and different subterranean habitats

as scree slopes and MSS7,11, limestone caves tend to be large and grow deep through dissolution of the rock by water during the genesis of the cave. The deeper the cave is, the lesser the

permeability of the rock, the epikarst usually reaches about 15 m deep24. In iron caves the fauna comes from the above ground through the MSS connections between surface and subterranean

environments, the same happens with energy that comes with roots that reach the MSS abundantly12. The troglobites develop in the MSS and eventually reach the caves where it can be seen in

its distribution limits, and the troglophiles go in the opposite direction, inhabiting the surface and going inside the caves to refuge from climate, but not going too far in the MSS (see

Table 1, species marked with1,3). PSEUDOCRYPTIC DIVERSITY Large caves with deep aphotic zones, stable abiotic conditions, water pools, often hosting bat colonies, are correlated to high

number of restricted species12, usually displaying classic troglomorphism as absence of eyes and body pigments, elongated appendages, increased body size25. In the ferruginous rock MSS the

same troglomorphisms are present in most species, even though, we observed that some Entomobryoid Collembola are often reduced in size, with normal or shortened (even though always

functional) appendages, similar to that of euedaphic fauna. Cryptic species recognized from a single widespread species complex through barcode sequencing, revealed related morphological

differences corresponding to the species separation26. To access this information it is necessary to expand the morphological refinement, some cryptic species are grouped together as result

of limited selection of diagnostic characters. This is by definition pseudocryptic species, when “individuals can be identified from morphology providing sufficient care is taken, but are so

similar that there is a high probability of misidentification, even by a competent scientist”27. The species found in ferruginous caves and MSS are very similar in most of its macro

morphology, differences are subtle, species recognition often must rely on minor details of chaetotaxy (Fig. 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24,

25, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 43, 44) and slight variations of morphological structures, often overlooked, as observed for the genera

_Arrhopalites_, _Pararrhopalites_, _Pseudosinella_ and _Trogolaphysa_. Such pseudocryptic diversity can only be accessed by specialized morphological scrutiny, molecular sequencing or a

combination of both. Whether we accept that cryptic diversity in Collembola cannot be explained by accelerated rates of molecular evolution28, it is likely that the diversity of subterranean

Collembola in ferruginous MSS and caves, results of the combination of the effects of lithology arrangement, phytophysiognomy and climate fluctuation at local scale. Finally, the

recognition of cryptic or pseudocryptic species within presumed widespread allopatric species is crucial to efficiently develop management and conservation plans22 and reduce the

underestimation of cave Collembola diversity. SUBTERRANEAN COLLEMBOLA HOTSPOT A total of 61 species of cave Collembola were recorded in Brazil so far, 40 are records from the State of Minas

Gerais, including 29 troglobites (Table 1), 26 of them are from iron caves and MSS as well as 11 troglophiles. For comparison we can consider two important subterranean hotspots in Limestone

caves in the States of Sao Paulo and Bahia (Southeastern and Northeastern regions), which presented an overall species richness of 28 and 22 troglobites, respectively. These two caves are

under different impact pressures, the former is in a protected area with controlled access, and the latter is under intense touristic exploitation3. Myers et al.1 combined richness,

endemism, distribution spam and threats to the area to define places of priority for conservation, called hotspots. The number of troglobites, with a full consideration of the threats or

conservation conditions of the caves and surroundings was, also, recently used as criteria for defining hotspot3,6. The ferruginous rock outcrops in Brazil are under a intense economic

pressure, the mining industry represents an important part of the production of commodities as iron ore and steel. The high diversity and endemism of cave Collembola found in recent studies

(Table 1), affecting directly the beta diversity of the areas considering the species are found nowhere else, and the continuous threat to the subterranean habitats formed in ferruginous

rock, justify categorizing the ferruginous subterranean habitats as hotspot for cave Collembola in the State of Minas Gerais. It is important to remark that the diversity considered here is

only for Collembola species, and that the studies mentioned above have a much higher phylogenetic diversity. CONSERVATION POLICY IMPLICATIONS The ferruginous caves and the MSS represent

sites of intense overlooked pseudocryptic diversification. Katz et al.22 observed that for Collembola in limestone areas the detection of short-range endemics, genetic isolation, and

apparent cryptic diversity has major conservation implications. The results we present here bring several considerations on conservation strategies and policies. The high diversity and

endemism rate observed for cave Collembola, associated to threats to the subterranean environments as mining, deforestation, and urbanization flag these areas as maximum priority and

interest for planning putative conservation areas29. These areas demand a multi-factor approach to successfully develop policies which optimize the diversity conservation, particularly

subterranean diversity. Brazilian legislation has protective measures for caves, but allows the complete suppression of a cave for mining or other exploratory purpose, under a process for

licensing the proposed activities. Even though some criteria are imposed, it fails in considering some important aspects of the cave structure in different lithology12. Under this

perspective the whole extension of ferruginous (and carbonatic) rock deposits in Brazil are available for exploitation, with irreversible impact on the subterranean fauna. There is over than

9400 companies in activity in the country, producing about 235.000.000 ton/year of iron ore, the second biggest production in the world. More than 72% of the Brazilian iron ore reservoirs

is located in the state of Minas Gerais, the locality of occurrence of 37 out of the 44 known species of Collembola found in ferruginous subterranean habitats in Brazil (Fig. 2). Here we

observed that the whole process needs a revision when comes to ferruginous rock, where the cave may not be the important spatial unit to preserve, instead, the high subterranean diversity

areas must be surveyed, not only in caves but also in the MSS. It is possible that in some cases to protect a hill that harbors a thick layer of _Canga_ with a troglobitic species rich MSS,

would result more effective to preserve restricted subterranean fauna, than to protect a small and shallow cave with reduced troglobitic richness. The state of Minas Gerais has 75 integral

conservation units (defined by law), with maximum protection policy, however, these conservation units represent only 1.05% (~ 619,800 ha) of the state territory. There are other categories

of conservation units, called of “sustainable use”, with much less restrictive policies. These categories of conservation units are much less effective to preserve epigean species, due to

the diverse usages and practices in those areas. Nevertheless, the 19 sustainable use conservation units in the state of Minas Gerais (private conservation units excluded) correspond to

3.01% (~ 1,768,000 ha) of the state territory (http://www.ief.mg.gov.br/unidades-de-conservacao—accessed Sep/31/2021). Sustainable use conservation units have some criteria that prevents

highly destructive activities, allowing some extractive crops, subsistence agriculture and tourism. These activities may be compatible with subterranean conservation through the MSS,

therefore the conservation unit network can get some advantage trying to connect integral conservation units with sustainable use ones. It was proposed that the sampling for subterranean

fauna in prospection drilling holes all over the area may bring important information about species richness and distribution, mainly if combined with cave and surface sampling30. This

procedure, implemented in the process for licensing new high impact exploratory activities, can improve the conservation effectiveness of the conservation units and compensation areas,

precisely define the role of the cave in the conservation plan, and shift the focus towards troglobitic species richness. CONCLUSIONS Our results depict the ferruginous subterranean

environment as an important hotspot for cave Collembola in the state of Minas Gerais, corroborating the expected high species richness in ferruginous rock caves and MSS. We also demonstrate

that access pseudocryptic diversity as observed in the genera _Arrhopalites_, _Pararrhopalites_, _Pseudosinella_ and _Trogolahysa_ is mandatory for planning the conservation strategies for

subterranean Collembola. The distribution of the species through the MSS can be favored by sustainable use conservation units, whether this fauna is surveyed along the licensing process.

Finally, we conclude that the conservation planning for future conservation unit establishment must focus not only on caves but also in the MSS, accessing the fauna through sampling in

prospection drilling holes. Protecting an area with high richness of endemic troglobites down in the MSS may be more effective than to protect a shallow cave when it comes to preserve

troglobitic diversity. METHODS PSEUDOCRYPTIC DIVERSITY The richness was the measure of the subterranean diversity, we surveyed all data about previous records for Brazilian Collembola cave

species, ecological status, lithology, and distribution from the literature, and included 11 newly found pseudocryptic species from subterranean habitats in iron and limestone rock. The

pseudocryptic species were verified by comparison of chaetotaxy and “micro-morphology” through optic and scanning microscopy of disjunct populations of a widespread morphotype. The imagery

was compared under hypotheses of chaetotaxic and morphologic homology, previously defined by different authors. Those populations with consistent discrete chaetotaxic and morphologic

patterns were assumed to be independent species, therefore they were taxonomically diagnosed, named, and ordered in a dichotomic identification key with all Brazilian species of the genus.

MICROSCOPY Specimens were preserved in ethanol 70% and mounted on slides following Jordana et al.31, after clearing using Nesbitt’s solution for study under phase contrast microscope, line

drawings were made with help of a drawing tube. For scanning electronic microscope (SEM) study, specimens were dehydrated by ethanol, dried in a critical point dryer, and covered in gold.

HOMOLOGY The terminology used in the diagnoses for the hypotheses of homology followed: labial chaetotaxy after Gisin32 with additions of Zhang and Pan33, Fjellberg34 for labial palp

papillae and maxillary palp; postlabial chaetotaxy after Chen and Christiansen35, with adaptations of Cipola et al.36 for J series; clypeal chaetotaxy after Yoshii and Suhardjono37; labral

chaetotaxy after Cipola et al.38; unguiculus lamellae after Hüther39; Anterior dens chaetotaxy after Oliveira et al.40; Mari-Mutt41 for dorsal head chaetotaxy, with additions of

Soto-Adames42; Szeptycki43 and Zhang and Deharveng44 for S-chaetotaxy; and Szeptycki45 for dorsal chaetotaxy, with additions and modifications provided by Soto-Adames42 and Zhang et al.46.

Symbols used to depict the chaetotaxy are presented in Fig. 4A–C. Codes will be used in _italics_ along the text to replace the morphological description of each chaeta and sensillum type.

Additional information about morphology and chaetotaxy of discussed species was obtained from the literature. ABBREVIATIONS USED IN THE DIAGNOSES Ant–antennal segment(s); b.c.–basal

chaeta(e), t.a.–terminal appendage of the maxillary palp; l.p.–lateral process of labial papilla E, lpc–labial proximal chaeta(e); Th–thoracic segment; Abd–abdominal segment(s);

Omt–trochanteral organ; a.e.–antero-external lamella, a.i.–antero-internal lamella, a.t.–unguis apical tooth, b.a.–basal anterior tooth of unguis, b.p.–basal posterior tooth of unguis,

m.t.–unguis median tooth, p.i.–postero-internal lamella, p.e.–postero-external lamella; mac–macrochaeta(e), mes–mesochaeta(e), mic–microchaeta(e), ms–specialized microchaeta(e),

psp–pseudopore(s), sens–specialized ordinary chaeta(e) (sensillum), MSS–Mesovoid Shallow Substratum. ECOLOGICAL STATUS To avoid subjectivity and ambiguity to determine the ecological status

of the species, we assumed to be a troglobite all the species with some degree of troglomorphism exclusively distributed in the subterranean environment, either caves, MSS, or both. Species

distributed in the surface and subterranean habitats were assumed to be troglophiles. IDENTIFICATION KEY FOR THE KNOWN AND NEW SPECIES OF THE GENUS _TROGOLAPHYSA_ RECORDED IN BRAZIL

TAXONOMIC DIAGNOSES AND MORPHOLOGICAL PLATES Type materials are deposited in the _Coleção de Referência de Fauna de Solo_, _Universidade Estadual da Paraíba_ (CRFS-UEPB) and _Museu Nacional

Rio de Janeiro, Universidade Federal do Rio de Janeiro_ (MNRJ-UFRJ). * https://zoobank.org:pub:2C8F4446-0869-48A9-ABD7-14D4C4DCB0FA * Additional records in Supplementary Material S1,

taxonomic references in S2. * Family Paronellidae Börner, 1906 * Subfamily Paronellinae Börner, 1906 * Tribe Paronellini sensu Zhang et al_._, 2019 * Genus _Trogolaphysa_ Mills, 1938 *

(Figs. 3, 4, 5, 6, 7, 8, 9, 10, 11) DIAGNOSIS Habitus typical of this genus (Fig. 3A–D), hyaline scales presents on Ant. I–II, head, body, and ventral face of furcula (Figs. 3C–D, 4A–C, 5D,

F, 7, 8, 11C), Ant IV smooth or annulated and never subdivided in two (Fig. 5A); eyes 0–8 (ex. Fig. 6C); prelabral and labral formula 4/5,5,4 (prelabral smooth or ciliate, pma smooth

chaetae) (Fig. 6A); antennobasal-organ present (Fig. 6C); labial chaetae L1–2 not reduced (Fig. 6E); sublobal plate of maxillary palp with 2 chaetae (Fig. 6E); Th II normally with A5 mac and

P3 complex with variable number of mac, and Th III with P3 mac present or abset (Fig. 7A, B), abdominal segments II–IV with 2, 3, 3 bothriotricha (Figs. 7C, D, 8A); unguis with three

external lamellae and unguiculus with P.E. lamella serrate or smooth (Fig. 9A–C); trochanteral organ with 2–4 psp (Fig. 10A) collophore anterior side with 2–3 distal mac (Fig. 10C);

tenaculum with four teeth on each branch and one anterior chaeta (Fig. 10D); manubrium without spines, manubrial plate with 2–3 PSP (Fig. 11A); anterior proximal dens with B.A., B.M. and I5

chaetae (Fig. 11B); dens with 1–2 rows of spines; mucro square or rectangular but relatively short, with 3–5 teeth (Fig. 11D). _Trogolaphysa bellinii_ SP. NOV. Oliveira, Lima & Zeppelini

Figures 12, 13 and 14, Tables 1 and 2 _Type material._ Holotype female in slide (15,482/CRFS-UEPB): Brazil, Minas Gerais State, Barão de Cocais municipality, cave MDIR-0028, next to “Mina

de Brucutu”, 19°52′48.7″S, 43°26′13.6″W, 19–23.viii.2019, Carste team coll. Paratypes in slides (15,468, 15,483/CRFS-UEPB): 2 females, same data as holotype. Paratypes in slides (15,519,

15,576/CRFS-UEPB donated to MNJR): 2 females, same data as holotype. Additional records see S1. _Description._ Total length (head + trunk) of specimens 1.53–1.75 mm (n = 5), holotype 1.70 

mm. Head. Ratio antennae: trunk = 1: 1.29–1.95 (n = 5), holotype = 1: 1.95; Ant III shorter than Ant II; Ant segments ratio as I: II, III, IV = 1: 1.80–2.24, 0.85–2.08, 0.85–2.08, holotype =

 1: 1.80, 0.85, 1.34. Antennal chaetotaxy: Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with a longitudinal row with about eight rod

sens, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without

modified sens, ventrally with one apical PSP, about three wrinkly sens on external longitudinal row, apical organ with two mic smooth chaetae externally, two coffee bean-like sens, and one

rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with four sub-apical finger-shaped sens, one wrinkly sens and two subapical rod sens,

ventrally with one apical PSP, about six wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three

basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0, rarely 2 + 2. Head dorsal chaetotaxy (Fig. 12A)

with 12 AN (AN1A–3), six A (A0–5), five M (M1–5), five S (S2–6), two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PA5 and PM3 as mes,

AN1A–3A with 10 mac plus two mes, A0 and A2 as mac; interocular P mes present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 12B).

Ventral chaetotaxy with 35–38 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of four to seven mes chaetae distally

(Fig. 12B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae

smooth (AN1–3, P2–3) and subequal in length (Fig. 12B). Maxillary palp with T.A. smooth and 1.23× larger than B.C. Thorax dorsal chaetotaxy (Fig. 13A). Th II A, M, P series with two mic

(A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with three mac, respectively, AL and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–7),

three mic (M4, M6–6P), three mes (M6E, M7–7E), four mic (P1–3, P6) respectively. Ratio Th II: III = 1.04–1.36: 1 (n = 5), holotype = 1.05: 1. Abdomen dorsal chaetotaxy (Fig. 13B, C). Abd I

A, M series with one (A5) and six (M2–6E) mic respectively, MS present. Abd II A, M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5

and M2 bothriotricha surrounded by five and four fan-shaped chaetae respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac

(M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae respectively; A5, M2 and M5 bothriotricha with six, two and three fan-shaped chaetae respectively, AS sens elongated, MS present.

Abd IV A–FE series with two mic (A1, A6), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, DE3), one mes

(D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), two mes (EE10–11), one mac (EE9), one mic (F1), two mes (F3, F3P), one mac (F2), one mic (FE2), three mes (FE3–5)

chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by five and two (T3) fan-shaped chaetae respectively; PS and AS present, and at least six supernumerary sens with uncertain

homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with four PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E), three mac (M2–3, M5), five mic

(P3A–6AE), one mic (P6E) two mes (AP6–PP6), four mac (P1, P3–5) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd III: IV = 1: 3.70–4.37 (n = 5), holotype = 1: 4.37. Legs. Trochanteral

organ diamond shape with about 20 spine-like chaetae, plus two PSP one external and one on distal vertex of Omt (Fig. 14A). Unguis outer side with one paired tooth straight and not developed

on proximal third; inner lamella wide with four teeth, basal pair subequal, B.P. not reaching the M.T. apex, M.T. just after the distal half, A.T. present. Unguiculus with lamellae smooth

and lanceolate (A.I., A.E., P.I.), except P.E. slightly serrate (Fig. 14B); ratio unguis: unguiculus = 1.56–1.79: 1 (n = 5), holotype = 1.56: 1. Tibiotarsal smooth chaetae about 0.9 × 

smaller than unguiculus; tenent hair capitate and about 0.55 × smaller than unguis outer lamella. Collophore (Fig. 14C). Anterior side with 12 ciliate, apically acuminate chaetae, five

proximal, four subdistal (as mes) and three distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row. Furcula. Covered with ciliate

chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 14D). Dens posterior face with two or more longitudinal rows of

spine-like chaetae about 24 external and 25 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.29 (holotype). _Etymology._

Species named after Dr. Bruno C. Bellini in recognition of his work on Brazilian Collembola. _Remarks. Trogolaphysa bellinii_ SP. NOV. resembles _T. bessoni_, _T. epitychia_ SP. NOV._,_ and

_T. mariecurieae_ SP. NOV. by 0 + 0 eyes (_T. bellinii_ SP. NOV. rarely with 2 + 2 eyes), Th II with 3 + 3 mac, and Th III without mac, but can be distinguished by presenting Abd IV with 4 +

 4 central mac (A3, A5, B4–5); _T. epitychia_ SP. NOV. with 3 + 3 central mac on Abd IV, _T. mariecuriea_e SP. NOV. with 2 + 2 central mac on Abd IV. _Trogolaphysa lacerta_ SP. NOV. Lima,

Oliveira & Zeppelini Figures 15, 16 and 17, Tables 1 and 2 _Type material._ Holotype male in slide (10,311/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Rio Acima municipality,

cave GAND-115, next to “Lapa do Calango”, 20°04′08.4″S, 43°40′09.9″W, 10.ii–20.iii.2014, Carste team coll. Paratypes in slides (10,312, 10,309/CRFS-UEPB): 2 males, same data as holotype.

Paratypes in slides (10,313, 10,314/CRFS-UEPB donated to MNJR): 2 females, same data as holotype. Additional records see S1. _Description._ Total length (head + trunk) of specimens 1.31–2.43

 mm (n = 5), holotype 1.86 mm. Head. Ratio antennae: trunk = 1: 1.33–1.46 (n = 2), holotype = 1: 1.46; Ant III shorter than Ant II; Ant segments ratio, I: II, III, IV = 1: 1.78–2.05:

1.5–1.64: 2.64–2.83, holotype = 1: 1.80: 1.64: 2.64. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens,

dorsally with a longitudinal row with about five rod sens, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate

mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP, one apical wrinkly sens on, apical organ with two coffee bean-like sens, and one rod sens

(Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with three sub-apical finger-shaped sens, one wrinkly sens and two apical rod sens, ventrally with

one apical PSP, one longitudinal external row with two subapical wrinkly sens and two medial finger-shaped sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and

mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0, rarely 3 + 3. Head

dorsal chaetotaxy (Fig. 15A) with 15 AN (AN1A–3), six A (A0–5), four M (M1–4), five S (S2–6), two PS (PS2, PS5), four PA (PA1–2, PA4–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4,

PE6) chaetae; PM3, PA5 and PP7 as mes, AN1A–3A with 11 mac plus four meso, A0 and A2 as mac; interocular P mes present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E

and L1–2 ciliate, R reduced (Fig. 15B). Ventral chaetotaxy with 36–38 ciliate chaetae and 1 reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row

of three to five mes chaetae distally (Fig. 15B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with L.P. finger-shaped and surpassing the base of

apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 15B). Maxillary palp with T.A. smooth and 1.28× larger than T.A. Thorax dorsal chaetotaxy (Fig. 

16A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with six mac, respectively, AL and MS present. Th III A, M, P series with

three mic (A1–3), two mes (A6–7), three mic (M4, M6–6P), three mes (M6E, M7–7E), four mic (P1–3, P6) respectively. Ratio Th II: III = 1.09–1.46: 1 (n = 5), holotype = 1.09: 1. Abdomen dorsal

chaetotaxy (Fig. 16B, C). Abd I M series with six (M2–6E) mic respectively, MS present. Abd II A, M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic

and AS present; A5 and M2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic

(M7I–7), three mac (M3, AM6, PM6), four mic (P6E, P7I–7P), one mac (P6) chaetae respectively; A5, M2 and M5 bothriotricha with seven, two and four fan-shaped chaetae respectively, AS sens

elongated, MS present. Abd IV A–FE series with four mic (A1, A5–6, AE1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), five mic (T1, T3, T5–7), five mic (D1–3,

DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), two mes (EE10–11), one mac (EE9), one mic (F1), two mes (F3–3P), one mac (F2), one mic (FE2), three mes

(FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and one fan-shaped chaetae respectively; PS and AS present, and at least six supernumerary sens with uncertain

homology _‘s’_(Fig. 8A); Abd. IV posteriorly with five to six PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E), three mac (M2–3, M5), five mic

(P3A–6AE), one mic (P6E) two mes (AP6–PP6), four mac (P1, P3–5) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd III: IV = 1: 3.70–4.37 (n = 5), holotype = 1: 4.37. Legs. Trochanteral

organ diamond shape with about 24 spine-like chaetae, plus two PSP one external and one on distal vertex of Omt (Fig. 17A). Unguis outer side with one paired tooth straight and not developed

on proximal third; inner lamella wide with four teeth, basal pair subequal, B.P. not reaching the M.T. apex, M.T. just after the distal half, A.T. present. Unguiculus with all lamellae

smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 17B); ratio unguis: unguiculus = 1: 1.50–1.79 (n = 5), holotype = 1: 1.75. Tibiotarsal smooth chaetae about 0.7× smaller than unguiculus;

tenent hair slightly acuminate and about 0.44× smaller than unguis outer lamella. Collophore (Fig. 17C). Anterior side with 10 ciliate, apically acuminate chaetae, five proximal (thinner);

three subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae

and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 17D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 50

external and 37 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.31 (n = 5). _Etymology. Lacerta_ from Latin means lizard, in

allusion to the name of the cave where this species was found, _Lapa do Calango_ (cave of the _Calango_), which is a small lizard common in this region. _Remarks. Trogolaphysa lacerta_ SP.

NOV. The new species resembles _T. caripensis, T. ernesti, T. piracurucaensis, T. formosensis_ and _T. dandarae_ SP. NOV. by the number of mac in Th II P3 complex (6 + 6), but is easily

distinguished by the head M2 and S5 mic (_T. caripensis_, _T. ernesti_, _T. formosensis_, _T. piracurucaensis_ as mac) and Th III without mac (_T. dandarae _SP. NOV. 3 + 3). _Trogolaphysa

chapelensis_ SP. NOV. Lima, Oliveira & Zeppelini Figures 18, 19 and 20, Tables 1 and 2 _Type material._ Holotype female in slide (4550/CRFS-UEPB): Brazil, Minas Gerais State, Rio Acima

municipality, cave Gruta-2d7, next to “Morro do Chapéu” 20°07′42.1″S, 43°54′26.2″W, 02–10.viii.2011, Andrade et al. coll. Paratypes in slides (4551–4553/CRFS-UEPB): 3 females, Brazil, Minas

Gerais State, Rio Acima municipality, cave Gruta-7d7, Qd7, 9d7 respectively, 20°07′42.1″S, 43°54′26.7″W, 29.iii–01.vi.2011, Andrade et al. coll. Paratype in slide (4603/CRFS-UEPB donated to

MNJR): 1 female, Brazil, Minas Gerais State, Rio Acima municipality, cave Gruta Qd7, 20°09′46.1″S, 43°49′36.2″W, 925 m, 29.iii–01.vi.2011, Andrade et al. Coll. Additional records see S1.

_Description._ Total length (head + trunk) 1.21–2.22 mm (n = 5), holotype 2.22 mm. Head. Ratio antennae: trunk = 1: 1.31–1.16 (n = 3), holotype = 1: 1.16; Ant III shorter than Ant II; Ant

segments ratio, I: II, III, IV = 1: 1.66–1.85, 1.65–1.78, 2.95–3.76, holotype = 1: 1.66, 1.65, 2.95. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short

ciliate mic and mac, and finger-shaped sens, dorsally with about six rod sens on longitudinal row, ventrally with one subapical-organ and about three subapical wrinkly sens (Fig. 4A); Ant

III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP, one apical wrinkly sens, apical

organ with two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about three sub-apical finger-shaped

sens and about three apical rod sens, ventrally with one apical PSP, one longitudinal external row with four wrinkly sens (Fig. 4A); and Ant I dorsally and ventrally with several short

ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head

dorsal chaetotaxy (Fig. 18A) with 15 AN (AN1A–3), six A (A0–5), four M (M1–4), five S (S2–6), two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6)

chaetae; PM3 and PA5 as mes, AN1A–3A with 13 mac plus two mes, A0 and A2 as mac; interocular P mic present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2

ciliate, R reduced (Fig. 18B). Ventral chaetotaxy with 29 ciliate chaetae; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of six mes chaetae distally (Fig. 18B).

Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth

(AN1–3, P2–3) and subequal in length (Fig. 18B). Maxillary palp with T.A. smooth and 1.17× larger than B.C. Thorax dorsal chaetotaxy (Fig. 19A). Th II A, M, P series with two mic (A1–2), one

mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with four mac, respectively, AL and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–7), two mic (M4–6P),

four mes (M6–6E, M7–7E), four mic (P1–3, P6) respectively. Ratio Th II: III = 1.10–1.31: 1 (n = 4), holotype = 1.10: 1. Abdomen dorsal chaetotaxy (Fig. 19B, C). Abd I A, M series with one

(A5) and six (M2–6E) mic respectively, MS present. Abd II A, M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha

surrounded by five and four fan-shaped chaetae respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three

mic (P6E, P7I–7), one mac (P6) chaetae respectively; A5, M2 and M5 bothriotricha with six, two and three fan-shaped chaetae respectively, AS sens elongated, MS present. Abd IV A–FE series

with three mic (A1, A6, AE1), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, DE3), one mes (D3P), one mic

(E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), two mes (EE10–11), one mac (EE9), one mic (F1), two mes (F3–3P), one mac (F2), one mic (FE2), three mes (FE3–5) chaetae,

respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae respectively; PS and AS present, and at least six supernumerary sens with uncertain homology

_‘s’_ (Fig. 8A); Abd. IV posteriorly with nine PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E), three mac (M2–3, M5), five mic (P3A–6AE), one

mic (P6E) two mes (AP6–PP6), four mac (P1, P3–5) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd III: IV = 1: 3.46–5.80 (n = 5), holotype = 1: 5.80. Legs. Trochanteral organ diamond

shape with about 23 spine-like chaetae, plus two PSP one external and one on distal vertex of Omt (Fig. 20A). Unguis outer side with one paired tooth straight and not developed on proximal

third; inner lamella wide with four teeth, basal pair subequal, B.P. not reaching the M.T. apex, M.T. just after the distal half, A.T. present. Unguiculus with lamellae smooth and lanceolate

(A.I., A.E., P.I.), except P.E. slightly serrate (Fig. 20B); ratio unguis: unguiculus = 1: 1.63–1.84 (n = 5), holotype = 1: 1.79. Tibiotarsal smooth chaetae about 0.8× smaller than

unguiculus; tenent hair capitate and about 0.52× smaller unguis outer lamella. Collophore (Fig. 20C). Anterior side with 13 ciliate, apically acuminate chaetae, seven proximal (thinner);

four subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae

and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 20D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 70 external

and 30 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.33 (n = 5). _Etymology._ Species named after Type locality _Morro do

Chapeu_. _Remarks. Trogolaphysa chapelensis_ SP. NOV. resembles _T. jacobyi_, _T. caripensis_, _T. bessoni_, and _T. belizeana_ by te absence of eyes (0 + 0 eyes) but is easily distinguished

by presenting 4 + 4 mac in Th II P3 complex (2–3 + 2–3 _T__. jacobyi_; 6 + 6 _T__. caripensis_; 2 + 2 _T__. bessoni_; 2–4 + 2–4 _T__. belizeana_), and 9 + 9 PSP posterior Abd IV (4 + 4T_.

belizeana_). _Trogolaphysa crystallensis_ SP. NOV. Oliveira, Lima & Zeppelini Figures 21, 22 and 23, Tables 1 and 2 _Type material._ Holotype female in slide (16,252/CRFS-UEPB): Brazil,

Minas Gerais State, Mariana municipality, cave LOC-0090, next to “Cachoeira Crystal”, 20°20′20.8″S, 43°23′44.3″W, 11–14.xi.2019, Carste team coll. Paratype in slide (16,251/CRFS-UEPB):

female, same data as holotype. Paratype in slide (16,254/CRFS-UEPB donated to MNJR): female, same data as holotype. Additional records see S1. _Description._ Total length (head + trunk) of

specimens 1.40–1.68 mm (n = 3), holotype 1.68 mm. Head. Ratio antennae: trunk = 1: 1.24–2.30 (n = 2), holotype = 1: 1.24; Ant III shorter than Ant II length; Ant segments ratio as I: II,

III, IV = 1: 1.72–1.78, 1.58–1.64, 3.11–3.14, holotype = 1: 1.78, 1.64, 3.14. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and

finger-shaped sens, dorsally with about three rod sens on longitudinal row, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several

short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP, about three wrinkly sens on external longitudinal row, apical organ with

two rod sens, and one finger-shaped sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with three sub-apical finger-shaped sens and one wrinkly

sens, ventrally with one apical PSP (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal

spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 21A) with 12–13 AN (AN1A–3), six A (A0–5), four M (M1–4), five S

(S2–6), two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PA5, PM3 and PP7 as mes, AN1A–3A, A0 and A2 as mac; interocular P mes present.

Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 21B). Ventral chaetotaxy with 33–35 ciliate chaetae and one reduced lateral spine;

postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of four to six mes chaetae distally (Fig. 21B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications.

Labial papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 21B). Maxillary palp with T.A.

smooth and 1.43 × larger than B.C. Thorax dorsal chaetotaxy (Fig. 22A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with

five mac, respectively, AL and MS present. Th III A, M, P series with two mic (A1–2), two mes (A6–7), theree mic (M4, M6–6P), three mes (M6E, M7–7E), four mic (P1–3, P6) respectively. Ratio

Th II: III = 1.05–1.27: 1 (n = 3), holotype = 1.05: 1. Abdomen dorsal chaetotaxy (Fig. 22B, C). Abd I A, M series with one (A5) and six (M2–6E) mic respectively, MS present. Abd II A, M, P

series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd

III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae respectively, A5, M2 and M5

bothriotricha with six, two and three fan-shaped chaetae respectively, AS sens elongated, MS present. Abd IV A–FE series with three mic (A1, A6, AE1), two mac (A3, A5), one mic (B1), one

mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–7), five mic (D1–1P, D3–3P, DE3), one mes (D2), two mes (E4P–4P2), three mac (E1–3), four mes (EE9–12), one mic (F1), three mes

(F2–3P), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by three and two (T3) fan-shaped chaetae respectively; PS and AS present, and at least

14 supernumerary sens with uncertain homology _‘s’_(Fig. 8A); Abd. IV posteriorly with three PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E),

three mac (M2–3, M5), five mic (P3A–P6AE), three mes (P6E–PP6), four mac (P1, P3–5) chaetae, respectively; AS and ACC.P4–5 present. Ratio Abd III: IV = 1: 4.06–4.51 (n = 3), holotype = 1:

4.51. Legs. Trochanteral organ diamond shape with about 18 spine-like chaetae, plus two PSP one external and one on distal vertex of Omt (Fig. 23A). Unguis outer side with one paired tooth

straight and not developed on proximal third; inner lamella wide with three teeth, basal pair subequal, B.P. little larger, but not reaching the M.T. apex, M.T. just after the distal half,

A.T. absent. Unguiculus with all lamellae smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 23B); ratio unguis: unguiculus = 1.48–1.79: 1 (n = 3), holotype = 1.48: 1. Tibiotarsal smooth

chaetae about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.5× smaller than unguis outer lamella. Collophore (Fig. 23C). Anterior side with 10 ciliate, apically acuminate

chaetae, six proximal, two subdistal (as mes) and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row. Furcula. Covered with

ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 23D). Dens posterior face with two or more longitudinal rows of

spine-like chaetae about 60 external and 28 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.31 (holotype). _Etymology._

Species named after Type locality _Cachoeira Crystal_ (Portuguese for Crystal falls). _Remarks. Trogolaphysa crystallensis_ SP. NOV. resembles _T. barroca_ SP. NOV., _T. gisbertae_ SP. NOV.,

_T. sotoadamesi_ SP. NOV., _T. triocelata_ and _T. zampauloi_ SP. NOV. by the absence of eyes (0 + 0 eyes) (_T. triocelata_ 3 + 3 and _T. zampauloi_ SP. NOV. eventually 4 + 4), Th II with 5

 + 5 mac, and Th III without mac. Can be distinguished from _T. barroca_ SP. NOV., _T. gisbertae_ SP. NOV., and _T. sotoadamesi_ SP. NOV. by the Abd IV with 4 + 4 central mac (A3, A5, B4–5);

_T. barroca_ SP. NOV., _T. gisbertae_ SP. NOV., and _T. triocelata_, with 3 + 3 and _T. sotoadamesi_ SP. NOV. 2 + 2 central mac on Abd IV. Finally, the new species differentiates from _T.

zampauloi_ SP. NOV. by unpaired lamella of unguis with one tooth, Omt with about 18 spine-like chaetae, dens external row with about 58 spines on _T. crystallensis_ SP. NOV. and unpaired

lamella of unguis with two teeth, Omt with about 26 spine-like chaetae, dens external row with about 30 spines on _T. zampauloi_ SP. NOV. _Trogolaphysa sotoadamesi_ SP. NOV. Ferreira,

Oliveira & Zeppelini Figures 24, 25 and 26, Tables 1 and 2 _Type material._ Holotype male in slide (13,162/CRFS-UEPB): Brazil, Minas Gerais State, Mariana municipality, cave ALEA 0003,

next to “Mina de Alegria”, 20°09′6.81″S, 43°29′13.6″W, 07.ii.2018, Bioespeloeo team coll. Paratypes in slides (13,146, 13,153/CRFS-UEPB): 2 females, same data as holotype, except 12.vi.2017.

Paratype in slide (13,173, 13,186/CRFS-UEPB donated to MNJR): 2 females, same data as holotype, except 09.vi.2017. Additional records see S1. _Description._ Total length (head + trunk) of

specimens1.50–1.81 mm (n = 5), holotype 1.50 mm. Head. Ratio antennae: trunk = 1: 1.26–1.45 (n = 3), holotype = 1: 1.38; Ant III shorter than Ant II; Ant segments ratio, I: II, III, IV = 1:

1.78–2.76, 1.52–2.22, 2.61–3.90, holotype = 1: 2.04, 1.68, 3.16. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped

sens, dorsally with a longitudinal row with about three rod sens, ventrally with one subapical-organ and with several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several

short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP, several wrinkly sens, apical organ with two coffee bean-like sens, one rod

sens and one finger-shaped sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with two sub-apical rod sens and two finger-shaped sens, ventrally

with one apical PSP and several finger-shaped sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally

with about seven basal spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 3A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 24A) with 16 AN (AN1A–3), six A (A0–5), four

M (M1–4), five S (S2–6), two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PM3 as mes (rarely mic), PA5 as mes, AN1A–3A, A0 and A2 as

mac; interocular P mes present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 24B). Ventral chaetotaxy with 37 ciliate chaetae and one

reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of six mes chaetae distally (Fig. 24B). Prelabral chaetae ciliate. Labral chaetae smooth, no

modifications. Labial papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–P3) and subequal in length (Fig. 24B).

Maxillary palp with T.A. smooth and 1.28× larger than B.C. Thorax dorsal chaetotaxy (Fig. 25A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic

(P4–6E), P3 complex with five mac, respectively, AL and MS present. Th III A, M, P series with three mic (A1–3, A6), one mes (A7), four mic (M4, M6–7, M6P), two mes (M6E, M7E), four mic

(P1–3, P6) respectively. Ratio Th II: III = 1.17–1.52: 1 (n = 5), holotype = 1.03: 1. Abdomen dorsal chaetotaxy (Fig. 25B, C). Abd I A, M series with one (A5) and six (M2–6E) mic

respectively, MS present. Abd II A, M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by five and

three fan-shaped chaetae respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one

mac (P6) chaetae respectively; A5, M2 and M5 bothriotricha with five, two and three fan-shaped chaetae respectively, AS sens elongated, MS present. Abd IV A–FE series with five mic (A1, A3,

A5–6, AE1), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), two mic (T1, T6), two mes (T5, T7), three mic (D1–2), two mes (D3P, DE3), two mes (E4P–P2), three mac (E1–3), one mic

(EE12), three mes (EE9–11), one mic (F1), two mes (F3–3P), one mac (F2), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and three

(T3) fan-shaped chaetae respectively; PS and AS present, and at least five supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with four PSP. Abd V A, M, P series

with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E), three mac (M2–3, M5), five mic (P3A–P6AE), one mic (P6E) two mes (AP6–PP6), four mac (P1, P3–5) chaetae, respectively;

AS, ACC.P4–5 present. Ratio Abd III: IV = 1: 5.03–4.42 (n = 5), holotype = 1: 4.42. Legs. Trochanteral organ triangular shape with about 19–21 spine-like chaetae, plus two PSP one external

and one on distal vertex of Omt (Fig. 26A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair unequal, B.P.

larger than B.A.; M.T. and A.T. absent. Unguiculus with all lamellae smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 26B); ratio unguis: unguiculus = 1: 1.46–1.91 (n = 5), holotype = 1:

1.91. Tibiotarsal smooth chaetae about 0.8 × smaller unguiculus; tenent hair acuminate and about 0.4 × smaller than unguis outer lamella. Collophore (Fig. 26C). Anterior side with seven

ciliate, apically acuminate chaetae, three proximal, two subdistal and two distal mac; lateral flap with nine chaetae, four ciliate in the proximal row and five smooth in the distal row.

Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (two inner mac) and three PSP (Fig. 26D). Dens posterior face with two or more

longitudinal rows of spine-like chaetae about 35 external and 26 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.39 (n = 5).

_Etymology._ Species named after Dr. Felipe N. Soto-Adames for his contribution on Collembola taxonomy and systematics. _Remarks. Trogolaphysa sotoadamesi_ SP. NOV. resembles _T. barroca_

SP. NOV., _T. crystallensis_ SP. NOV., _T. gisbertae_ SP. NOV., _T. zampauloi_ SP. NOV. by 0 + 0 eyes (_T. zampauloi_ SP. NOV. rarely with 4 + 4 eyes), Th II P3 complex with five mac, Th III

without mac, manubrial plate with five ciliate chaetae and mucro with four teeth. The new species _T. sotoadamesi_ SP. NOV. with 2 + 2 central mac on Abd IV differentiates from _T. barroca_

SP. NOV., _T. gisbertae_ SP. NOV. with 3 + 3, and _T. crystallensis_ SP. NOV., _T. zampauloi_ SP. NOV. with 4 + 4 central mac. _Trogolaphysa mariecurieae_ SP. NOV. Ferreira, Oliveira &

Zeppelini Figures 27, 28 and 29, Tables 1 and 2 _Type material._ Holotype female in slide (9109/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Mato Dentro municipality, MSS 10/11, next

to “Pico do Soldado” 19°00′23.86″S, 43°23′41.266″W, 11–10.ix.2015, Carste team coll. Paratypes in slides (5888, 5857/CRFS-UEPB): 2 females, same data as holotype, except,19–23.v.2014,

Soares et al_._ coll. Paratype in slide (9222, 10,760/CRFS-UEPB donated to MNJR): 2 females, same data as holotype, except 19°00′18.72″S, 43°23′30.031″W, 14.x.2015 and 18–20.iv.2016.

Additional records see S1. _Description._ Total length (head + trunk) of specimens 1.07–1.49 mm (n = 5), holotype 1.49 mm. Head. Ratio antennae: trunk = 1: 1.69–1.91 (n = 2), holotype = 1:

1.69; Ant III shorter than Ant II length; Ant segments ratio, I: II, III, IV = 1: 2.00–2.75, 1.69–2.55, 4.02–5.29, holotype = 1: 2.75, 1.69, 4.02. Antennal chaetotaxy (no represented): Ant

IV dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally with one longitudinal row with about four rod sens, ventrally with one subapical-organ

and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one

apical PSP, apical organ with two rod sens (Fig. 4A); Ant II dorsally and ventrally with several short less ciliate mic and mac, dorsally with five apical rod sens, ventrally with one apical

PSP, about five wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short less ciliate mic and mac, dorsally with three basal spine-like sens,

ventrally with seven basal spine-like sens, about five smooth mic, and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 27A) with 12 AN (AN1A–3), six A (A0–5),

four M (M1–4), five S (S2–6), two PS (PS2, PS5), four PA (PA1–3, PA5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PM3 and PA5 as mes, AN1A–3A, A0 and A2 as mac;

interocular P mic present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 27B). Ventral chaetotaxy with 34 ciliate chaetae and one

reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of six mes chaetae distally (Fig. 27B). Prelabral chaetae ciliate. Labral chaetae smooth, no

modifications. Labial papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 27B). Maxillary

palp with T.A. smooth and 1.13× larger than B.C. Thorax dorsal chaetotaxy (Fig. 28A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3

complex with three mac, respectively, AL and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–7), three mic (M4–M6P), three mes (M6E, M7–7E), four mic (P1–3, P6)

respectively. Ratio Th II: III = 0.85–1.02: 1 (n = 4), holotype = 0.89: 1. Abdomen dorsal chaetotaxy (Fig. 28B, C). Abd I A, M series with one (A5) and six (M2–6E) mic respectively, MS

present. Abd II A, M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and two fan-shaped

chaetae respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), two mic (P6E, P7I), one mac (P6) chaetae

respectively; A5, M2 and M5 bothriotricha with five, two and two fan-shaped chaetae respectively, AS sens elongated, MS present. Abd IV A–FE series with three mic (A1, A6, AE1), one mac

(A4), two mic (B1–2), one mes (B6), one mac (B5), four mic (C1–4), three mic (T1, T5, T7), five mic (D1–3, DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic

(EE12), two mes (EE10–11), one mac (EE9), one mic (F1), three mes (F2–3P), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and three

(T3) fan-shaped chaetae respectively; PS and AS present, and at least five supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with four PSP. Abd V A, M, P series

with two mic (A1, A3), one mes (A6), one mac (A5), five mac (M2–3, M5–5E), five mic (P3A–P6AE), two mes (AP6–PP6), four mac (P1, P3–5) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd

III: IV = 1: 4.27–5.91 (n = 5), holotype = 1: 5.02. Legs. Trochanteral organ diamond shape with about 15 spine-like chaetae, plus 2–3 PSP one external, one on distal vertex and another

(present or absent) on top of posterior spines row of Omt (Fig. 29A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth,

basal pair subequal, B.P. larger than B.A., inner lamella with unpaired small M.T. between B.A. and B.P. and A.T. absent. Unguiculus with all lamellae smooth and truncate (A.I., A.E., P.I.,

P.E.) (Fig. 29B); ratio unguis: unguiculus = 1.50–1.95: 1 (n = 5), holotype = 1.95: 1. Tibiotarsal smooth chaetae about 0.9× smaller than unguiculus; tenent hair slightly capitate and about

0.6× smaller than unguis outer lamella. Collophore (Fig. 29C). Anterior side with eight ciliate, apically acuminate chaetae, six proximal and two distal mac; lateral flap with 13 chaetae,

five ciliate in the proximal row and eight smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two

inner mac) and three PSP (Fig. 29D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 40 external and 22 internal, external spines larger and thinner than

internal ones. Mucro with four teeth, ratio width: length = 0.23 (holotype). _Etymology._ Species named after Dr. Marie Skłodowska-Curie for her enormous contribution to science. _Remarks.

Trogolaphysa mariecurieae_ SP. NOV. resembles _T. bellinii_ SP. NOV. _T. jacobyia_ and _T. epitychia_ SP. NOV. by the absence of eyes (_T. bellinii_ SP. NOV. rarely with 2 + 2 eyes), Th II

P3 complex with three mac and with one unpaired tooth on inner lamella of unguis. The new species _T. mariecurieae_ SP. NOV. (Abd IV with 2 + 2 mac) differs from _T. jacobyia, T. epitychia_

SP. NOV. both with Abd IV 3 + 3, and _T. bellinii_ SP. NOV. with 4 + 4 central mac. _T. mariecurieae_ SP. NOV. and _T. bellinii_ SP. NOV. with capitate tenent hair, in contrast with _T.

jacobyia_ and _T. epitychia _SP. NOV_._ with acuminated tenant hair. _Trogolaphysa barroca_ SP. NOV. Brito & Zeppelini Figures 30, 31 and 32, Tables 1 and 2 _Type material._ Holotype

female in slide (13,167/CRFS-UEPB): Brazil, Minas Gerais State, Mariana municipality, ALFA-0003 cave, 20°09′06.8″S, 43°29′13.6″W, 07–27.ii.2018, Bioespeleo team coll. Paratype in slide

(13,150/CRFS-UEPB): 1 female, same data as holotype, except 12.vi.2017. Paratype in slide (13,158/CRFS-UEPB donated to MNJR): 1 female, same data as holotype. Paratype in slide

(13,197/CRFS-UEPB): 1 female, Brazil, Minas Gerais State, Mariana municipality, ALEA-0004 cave, 20°09′00.0″S, 43°29′11.8″W, 07.ii.2018, Bioespeleo team coll. Paratype in slide

(13,203/CRFS-UEPB): 1 female, Brazil, Minas Gerais State, Mariana municipality, ALEA-0002 cave, 20°08′56.5″S, 43°29′09.8″W, 27.ii.2018, Bioespeleo team coll. Additional records see S1.

_Description._ Total length (head + trunk) of specimens 1.70–2.13 mm (n = 5), holotype 1.81 mm. Head. Ratio antennae: trunk = 1: 1.27–1.60 (n = 3), holotype = 1: 1.27; Ant III shorter than

Ant II; Ant segments ratio as, I: II, III, IV = 1: 1.90–2.41, 1.64–2.02, 2.69–3.64, holotype = 1: 1.90, 1.67, 2.69. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with

several short less ciliate mic and mac, and finger-shaped sens, dorsally with about four rod sens on longitudinal row, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A);

Ant III dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP, about nine wrinkly sens on

external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short less ciliate mic

and mac, dorsally with two sub-apical finger-shaped sens and two subapical rod sens, ventrally with one apical PSP, and several wrinkly sens on longitudinal external row (Fig. 4A); and Ant I

dorsally and ventrally with several short less ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with about five basal spine-like sens, about five smooth mic and

several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 30A) with 14–15 AN (AN1A–3), six A (A0–5), five M (M1–5), six S (S1–6), two PS (PS2, PS5), four PA (PA1–3,

PA5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PM3 as mic, A3 as mes, AN1A–3, A0, A2 and PA5 as mac; interocular P mic present. Basomedian and basolateral labial

fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 30B). Ventral chaetotaxy with 33 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2,

chaetae B.C. present and a collar row of five to six mes chaetae distally (Fig. 30B). Prelabral chaetae weakly ciliate. Labral chaetae smooth, no modifications. Labial papilla E with L.P.

finger-shaped and subequal the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3), and subequal in length (Fig. 30B). Maxillary palp with T.A. smooth and 1.14 × larger

than B.C. Thorax dorsal chaetotaxy (Fig. 31A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with five mac, respectively, AL

and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–7), two mic (M4, M6P), four mes (M6–6E, M7–7E), and four mic (P1–3, P6), respectively. Ratio Th II: III = 1.11–1.35:

1 (n = 5), holotype = 1.29: 1. Abdomen dorsal chaetotaxy (Fig. 31B, C). Abd I A, M series with one (A5) and six (M2–6E) mic, respectively, MS present. Abd II A, M, P series with two mic

(A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and three fan-shaped chaetae, respectively. Abd III A, M, P series

with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae, respectively; A5, M2 and M5 bothriotricha

with six, two and three fan-shaped chaetae, respectively; AS sens elongated, MS present. Abd IV A–FE series with four mic (A1, A5–6, AE1), one mac (A3), one mic (B1), one mes (B6), two mac

(B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), three mes (EE9–11), one mic

(F1), three mes (F2–3P), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae, respectively; PS and AS

present, and at least seven supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with four to six PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6),

one mac (A5), two mes (M5A–5E), three mac (M2–3, M5), five mic (P3A–6AE), one mic (P6E), two mes (AP6, PP6), four mac (P1, P3–5) chaetae, respectively; AS and ACC.P4–5 present. Ratio Abd

III: IV = 1: 3.38–5.55 (n = 5), holotype = 1: 5.27. Legs. Trochanteral organ diamond shape with about 16–21 spine-like chaetae, plus 2–3 PSP one external, and two (one of them present or

absent) on top of posterior spines row of Omt (Fig. 32A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair

subequal; B.P. little larger than B.A., M.T. and A.T. absent. Unguiculus with all lamellae smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 32B); ratio unguis: unguiculus = 1.53–1.67: 1

(n = 5), holotype = 1.61: 1. Tibiotarsal smooth chaetae about 0.61 × smaller than unguiculus; tenent hair acuminate and about 0.4 × smaller than unguis outer lamella. Collophore (Fig. 32C).

Anterior side with eight ciliate, apically acuminate chaetae, four proximal (thinner), one subdistal and three distal mac; lateral flap with 10 chaetae, five ciliate in the proximal row and

five smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (three inner mac) and three PSP (Fig. 32D).

Dens posterior face with two or more longitudinal rows of spines-like chaetae about 22 external and 37–39 internal, external spines larger and thinner than internal ones. Mucro with four

teeth, ratio width: length = 0.33 (holotype). _Etymology._ Refers to the Baroque art (which is “barroco” noun, in Portuguese) of Mariana, Minas Gerais, type locality. _Remarks. Trogolaphysa

barroca_ SP. NOV. resembles _T. formosensis_ by head PM3 mic (mac in _T. piracurucaensis_, _T. gisbertae_ SP. NOV. and _T. dandarae_ SP. NOV.; mes in _T. ernesti_, _T. sotoadamesi_ SP. NOV.

and _T. mariecurieae_ SP. NOV.); 3 + 3 head dorsal mac like _T. ernesti_, although in the new species it is as A0, A2 and PA5, and in _T. ernesti_ is A0, A2–3; unguis M.T. and A.T. teeth

absent like _T. sotoadamesi_ SP. NOV. and _T. dandarae_ SP. NOV. (present in _T. bellini_ SP. NOV., _T. lacerta_ SP. NOV. and _T. chapelensis_ SP. NOV.). _Trogolaphysa epitychia_ SP. NOV.

Oliveira, Lima & Zeppelini Figures 33, 34 and 35, Tables 1 and 2 _Type material._ Holotype male in slide (10,578/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Mato Dentro

municipality, cave CSS-0118, next to “São Sebastião do Bom Sucesso”, 18°56′14.1″S, 43°24′43.8″W, 21.xi–15.xii.2016, Carste team coll. Paratypes in slides (10,580, 10,585/CRFS-UEPB): 2

females, same data as holotype. Paratypes in slides (10,653, 10,692/CRFS-UEPB donated to MNJR): 1 female and 1 male, same data as holotype, except 22.xi–15.xii.2016 and 31.v–12.vi.2016,

respectively. Additional records see S1. _Description._ Total length (head + trunk) 1.13–1.35 mm (n = 5), holotype 1.13 mm. Head. Ratio antennae: trunk = 1: 1.29–1.95 (n = 5), holotype = 1:

1.95; Ant III shorter than Ant II; Ant segments ratio as I: II, III, IV = 1: 1.69–2.20, 1.14–1.86, 2.37–3.52, holotype = 1: 1.71, 1.14, 2.37. Antennal chaetotaxy (no represented): Ant IV

dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with one longitudinal row with about six rod sens, ventrally with one subapical-organ and one

longitudinal row with about four wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens,

ventrally with one apical PSP, about three wrinkly sens on external longitudinal row, apical organ with two coffee bean-like sens, one rod sens and one smooth mic (Fig. 4A); Ant II dorsally

and ventrally with several short ciliate mic and mac, dorsally with about six sub-apical finger-shaped sens and one wrinkly sens, ventrally with one apical PSP, about three wrinkly sens on

longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, three basal spine-like sens, ventrally with four basal spine-like sens, about

three smooth mic, several finger-shaped sens, and two wrinkly sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 33A) with 12 AN (AN1A–3), six A (A0–5), four M (M1–4), five S (S2–6),

two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PM3 and PA5 as mes, AN1A–3A, A0 and A2 as mac; interocular P mes present. Basomedian

and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 33B). Ventral chaetotaxy with 31–32 ciliate chaetae and one reduced lateral spine; postlabial G1–4;

X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of five to six mes chaetae distally (Fig. 33B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E

with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 33B). Maxillary palp with T.A. smooth and

1.26× larger than B.C. Thorax dorsal chaetotaxy (Fig. 34A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with three mac,

respectively, AL and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–A7), three mic (M4, M6–6P), three mes (M6E, M7–7E), four mic (P1–3, P6) respectively. Ratio Th II:

III = 1.05–1.21: 1 (n = 5), holotype = 1.18: 1. Abdomen dorsal chaetotaxy (Fig. 34B, C). Abd I A, M series with one (A5) and six (M2–6E) mic respectively, MS present. Abd II A, M, P series

with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III A, M,

P series with two mic (A7I–7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae, respectively; A5, M2 and M5

bothriotricha with five, two and one fan-shaped chaetae, respectively; AS sens elongated, MS present. Abd IV A–FE series with three mic (A1, A6, AE1), two mac (A3, A5), two mic (B1, B4), one

mes (B6), one mac (B5), four mic (C1–4), four mic (T1, T3, T5–6), one mac (T7), six mic (D1–3P, DE3), two mic (E4P–4P2), three mac (E1–3), one mic (EE11), three mes (EE9–10, EE12), one mic

(F1), three mes (F2–3P), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by five and two fan-shaped chaetae, respectively; PS and AS present,

and at least seven supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with three PSP. Abd V A, M, P series with three mic (A1, A3, A6), one mac (A5), two mic

(M3, ME5), three mac (M2, M5–5A), two mic (P3A–4A), one mes (P5A) two mac (P6AI–6AE), four mes (P5–PP6), three mac (P1, P3–4) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd III: IV =

 1: 4.69–5.55 (n = 5), holotype = 1: 4.88. Legs. Trochanteral organ in V–shape with about 15 spine-like chaetae, plus 4 PSP one external, one on distal vertex and another two on top of

posterior spines row of Omt (Fig. 35A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, B.P.

little larger, not reaching the M.T. apex, M.T. just after the distal half, A.T. absent. Unguiculus with all lamellae smooth and slightly truncate (A.I., A.E., P.I., P.E.) (Fig. 35B); ratio

unguis: unguiculus = 1.17–1.98: 1 (n = 5), holotype = 1.17: 1. Tibiotarsal smooth chaetae about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.53× smaller than unguis outer

lamella. Collophore (Fig. 35C). Anterior side with nine ciliate, apically acuminate chaetae, five proximal, two subdistal and two distal mac; lateral flap with 10 chaetae, five ciliate in

the proximal row and five smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (two inner mac) and three

PSP (Fig. 35D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 60 external and 34 internal, external spines larger and thinner than internal ones. Mucro

with four teeth, ratio width: length = 0.30 (holotype). _Etymology. Epitychia_ from Greek means success, in allusion to the collection site where the species was found _São Sebastião do Bom

Sucesso_. _Remarks. Trogolaphysa epitychia_ SP. NOV. resembles _T. bellinii_ SP. NOV., _T. bessoni,_ and _T. mariecurieae_ SP. NOV. by the absence of eyes (_T. bellinii_ SP. NOV. rarely with

2 + 2 eyes), Th II with 3 + 3 mac, and Th III without mac. Differentiates from _T. bellinii_ SP. NOV. and _T. mariecurieae_ SP. NOV. by Abd IV with 3 + 3 (A3, A5, B5), 4 + 4, and 2 + 2 mac

on Abd IV respectively; on _T. bellinii_ SP. NOV. and can be distinguished from _T. bessoni_ by the absence of unpaired tooth on inner lamella of unguis, external row of dens with 25 spines,

inner row of dens with 20 spines (_T. epitychia _SP. NOV. with one unpaired tooth M.T. on inner lamella of unguis, external row of dens with about 60 spines and inner row of dens with about

34 spines). _Trogolaphysa zampauloi_ SP. NOV. Lima, Oliveira & Zeppelini Figures 36, 37 and 38, Tables 1 and 2 _Type material._ Holotype female in slide (11,851/CRFS-UEPB): Brazil, São

Paulo State, Ribeira municipality, cave MTD-13, nexto to “Serra Pontalhão”, 24°38′47.4″S, 48°57′52.6″W, 08–20.iii.2016, Carste team coll. Paratypes in slides (11,875–11,878/CRFS-UEPB): 2

males and 2 females, Brazil, São Paulo State, Ribeira municipality, cave MTD-02, 24°37′27.3″S, 48°57′35.8″W, 08–20.iii.2016. Paratype in slide (11,876/CRFS-UEPB donated to MNJR). Additional

records see S1. _Description._ Total length (head + trunk) of specimens 1.35–1.91 mm (n = 5), holotype 1.35 mm. Head. Ratio antennae: trunk = 1: 1.35–1.55 (n = 2), holotype = 1: 1.55; Ant

III smaller than Ant II length; Ant segments ratio as I: II, III, IV = 1: 1.71–2.38, 1.60–1.88, 2.85–3.61, holotype = 1: 2.38, 1.88, 3.61. Antennal chaetotaxy (no represented): Ant IV

dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about three rod sens on longitudinal row, ventrally with one subapical-organ, and about

three wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical PSP,

one apical wrinkly sens, apical organ with two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about

three sub-apical finger-shaped sens and two apical rod sens, ventrally with one apical PSP, one longitudinal external row with two subapical finger-shaped sens and two medial wrinkly sens

(Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about four smooth

mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0 to 4 + 4. Head dorsal chaetotaxy (Fig. 36A) with 14 AN (AN1A–3), six A (A0–5), four M (M1–4), five S (S2–6), two PS (PS2, PS5), four

PA (PA1–3, PA5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; PE4, PE6, PM3 and PA5 as mes, AN1A–3A as mac, A0 and A2 as mac, A3–5 as mes; interocular P mes present.

Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 36B). Ventral chaetotaxy with about 37 ciliate chaetae, plus one reduced lateral spine;

postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of eight mes chaetae distally (Fig. 36B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial

papilla E with L.P. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 36B). Maxillary palp with T.A.

smooth and 1.17 × smaller than B.A. Thorax dorsal chaetotaxy (Fig. 37A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with

five mac, respectively, AL and MS present. Th III A, M, P series with three mic (A1–3), two mes (A6–A7), three mic (M4, M6–6P), three mes (M6E, M7–7E), four mic (P1–3, P6), respectively.

Ratio Th II: III = 1.02–1.48: 1 (n = 5), holotype = 1.21: 1 Abdomen dorsal chaetotaxy (Fig. 37B, C). Abd I A, M series with one (A5) and six (M2–6E) mic respectively, MS present. Abd II A,

M, P series with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL as mic and AS present; A5 and M2 bothriotricha surrounded by three and two fan-shaped chaetae

respectively. Abd III A, M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae

respectively; A5, M2 and M5 bothriotricha with five, two and three fan-shaped chaetae respectively, AS sens elongated, MS present. Abd IV A–FE series with three mic (A1, A6, AE1), two mac

(A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac

(E1–3), one mic (EE12), one mes (EE11), two mac (EE9–10), one mic (F1), two mes (F3–3P), one mac (F2), one mic (FE2), two mes (FE3, FE5), one mac (FE4) chaetae, respectively; T2, T4 and E4

bothriotricha surrounded by four and four (T3) fan-shaped chaetae respectively; PS and AS present, and at least six supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV

posteriorly with three PSP. Abd V A, M, P series with two mic (A1, A3), one mes (A6), one mac (A5), two mes (M5A, M5E), three mac (M2–3, M5), five mic (P3A–6AE), one mic (P6E) two mes

(AP6–PP6), four mac (P1, P3–5) chaetae, respectively; AS, ACC.P4–5 present. Ratio Abd III: IV = 1: 3.29–4.28 (n = 5), holotype = 1: 4.10. Legs. Trochanteral organ diamond shape with about 27

spine-like chaetae, plus 3–4 PSP one external, one on distal vertex and another two (one of them present or absent) on top of posterior spines row of Omt (Fig. 38A). Unguis outer side with 

one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, B.P. not reaching the M.T. apex, M.T. just after the distal half, A.T.

 present. Unguiculus with all lamellae smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 38B); ratio unguis: unguiculus = 1.63–1.84 (n = 5), holotype = 1: 1.79. Tibiotarsal smooth chaetae

about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.39× smaller than unguis outer edge. Collophore (Fig. 38C). Anterior side with five ciliate, apically acuminate chaetae,

 two proximal (thinner); one subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row. Furcula. Covered with ciliate

chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 38D). Dens posterior face with two or more longitudinal rows of

spine-like chaetae about 30 external and 23 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.29 (n = 5). Etymology. Species

named after the field biologist MSc. Robson de Almeida Zampaulo for his contribution to Brazilian biospeleology. Remarks. _Trogolaphysa zampauloi_ SP. NOV. resembles _T. caripensis; T.

ernesti; T. piracurucaensis_ by Th III without mac, and 4 + 4 central mac (A3, A5, B4–5) in Abd IV, but is easily distinguished from these species by the presence of Th II with 4 + 4 mac in

P3 complex (6 + 6T_. caripensis, T. ernesti, T. piracurucaensis_). For more comparisons see remarks in _T. crystallensis_ SP. NOV. _Trogolaphysa gisbertae_ SP. NOV. Brito & Zeppelini

Figures 39, 40 and 41, Tables 1 and 2 _Type material._ Holotype female in slide (6668/CRFS-UEPB): Brazil, Pará State, Parauapebas municipality, cave N1N8-N8-017, next to “Serra Norte”,

06°10′05.9″S, 50°09′25.6″W, 02–29.iv.2015, Carste team coll. Paratype in slide (6669/CRFS-UEPB donated to MNJR): 1 female, same data as holotype, except 04.ix–06.x.2014. Paratype in slide

(6973/CRFS-UEPB): 1 female, same data as holotype, except 04.ix–06.x.2014. Paratypes in slides (6657, 7138/CRFS-UEPB): 2 females, Brazil, Pará State, Parauapebas municipality, N1N8-N8-020

cave, 06°10′07.8″S, 50°09′25.4″W, 17.vii–04.viii.2014, Carste team coll. Additional records see S1. _Description._ Total length (head + trunk) of specimens 1.10–1.23 mm (n = 5), holotype

1.15 mm. Head. Ratio antennae: trunk = 1: 1.44–1.55 (n = 3); Ant segments ratio as I: II, III, IV = 1: 1.67–2.43, 1.58–2.63, 2.91–5.46, holotype = 1: 2.03, – , 3.90. Antennal chaetotaxy (no

represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about five rod sens in row, ventrally with one subapical-organ and

several wrinkly sens row (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical

PSP, about four wrinkly sens on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with

several short ciliate mic and mac, dorsally with four finger-shapedd sens in row and two subapical rod sens, ventrally with one apical PSP, and about five wrinkly sens on longitudinal

external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about

five smooth mic and several fniger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 39A) with 11 AN (AN1A–3), six A (A0–5), four M (M1–4), five S (S1–5), two PS (PS2, PS5),

four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; AN1A–3A, A0, A2–3, PA5 and PM3 as mac; interocular P absent. Basomedian and basolateral labial fields

with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 39B). Ventral chaetotaxy with 20 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C.

present and a collar row of three to four mes chaetae distally (Fig. 39B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with L.P. finger-shaped and

surpassing the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 39B). Maxillary palp with T.A. smooth and 1.32 × larger than B.C. Thorax

dorsal chaetotaxy (Fig. 40A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with five mac, respectively, AL and MS presents.

Th III A, M, P series with three mic (A1–3), two mes (A6–7), three mic (M4, M6–6P), three mes (M6E, M7–7E), and four mic (P1–3, P6), respectively. Ratio Th II: III = 1.00–2.60: 1 (n = 5),

holotype = 1.28: 1. Abdomen dorsal chaetotaxy (Fig. 40B, C). Abd I A, M series with one (A5) and six (M2–6E) mic, respectively, MS present. Abd II A, M, P series with two mic (A6–7), two mac

(M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and two fan-shaped chaetae, respectively. Abd III A, M, P series with one mic

(A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6), three mic (P6E, P7I–7), one mac (P6) chaetae, respectively; A5, M2 and M5 bothriotricha with six, two

and three fan-shaped chaetae, respectively, AS sens elongated, MS present. Abd IV A–FE series with four mic (A1, A5–6, AE1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four

mic (C1–4), four mic (T1, T5–7), five mic (D1–3, DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), three mes (EE9–11), one mic (F1), three mes (F2–3P),

one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae, respectively; PS and AS present, and at least six

supernumerary sens with uncertain homology _‘s’ _(Fig. 8A); Abd. IV posteriorly with one to three PSP. Abd V A, M, P series with three mic (A1, A3), one mes (A6), one mac (A5), two mes

(M5A–5E), three mac (M2–3, M5), five mic (P3A–6AE), one mic (P6E), two mes (AP6, PP6), four mac (P1, P3–5) chaetae, respectively; AS and ACC.P4–5 present. Ratio Abd III: IV = 1: 3.29–4.90 (n

 = 5), holotype = 1: 3.29. Legs. Trochanteral organ diamond shape with about 25 spine-like chaetae, plus two PSP one external, and one on distal vertex of Omt (Fig. 41A). Unguis outer side

with one paired tooth straight and not developed on proximal third; inner lamella wide with three teeth, basal pair subequal, B.P. not reaching the M.T. apex, M.T. just after the distal

half, A.T. absent. Unguiculus with lamellae smooth and slightly truncate (A.I., A.E., P.I.), except P.E. slightly serrate (Fig. 41B); ratio unguis: unguiculus = 1.59–2.05: 1 (n = 5),

holotype = 1.62: 1. Tibiotarsal smooth chaetae about 0.9× smaller than unguiculus; tenent hair acuminate and about 0.53× smaller than unguis outer lamella. Collophore (Fig. 41C). Anterior

side with five ciliate, apically acuminate chaetae, one proximal (thinner); two subdistal and two distal mac; lateral flap with 10 chaetae, five ciliate in the proximal row and five smooth

in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (three inner mac) and three PSP (Fig. 41D). Dens posterior

face with two or more longitudinal rows of spine-like chaetae about 18 external and 24 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width:

length = 0.26 (holotype). _Etymology._ Honor to Gisberta Salce Júnior, Brazilian woman, murdered in 2006 (Porto, Portugal) in a transphobia crime. _Remarks. Trogolaphysa gisbertae_ SP. NOV.

differs from _T. ernesti_ and _T. formosensis_ (with 0 + 0 head dorsal mac), T_. piracurucaensis_, and _T. barroca_ SP. NOV. (1+1 head dorsal mac); and resembles _T. dandarae_ SP. NOV. (with

5+5 head dorsal mac), but it is easily distinguishable by Th II P3 complex and Th III mac (5 + 5 and 0 + 0, 6 + 6 and 3 + 3, respectively); and unguis with M.T. present (absent in _T.

sotoadamesi_ SP. NOV., _T. barroca_ SP. NOV.). _Trogolaphysa dandarae_ SP. NOV. Brito & Zeppelini Figures 42, 43 and 44, Tables 1 and 2 _Type material._ Holotype female in slide

(12,775/CRFS-UEPB): Brazil, Pará State, Parauapebas municipality, cave N4WS-0018/48, next to “Serra Norte”, 06°04′34.5″S, 50°11′37.7″W, 21–30.vii.2018, Brandt Meio Ambiente team coll.

Paratype in slide (12,776/CRFS-UEPB donated to MNJR): 1 female, same data as holotype. Paratypes in slides (12,777, 12,778/CRFS-UEPB): 2 females, same data as holotype. Paratypes in slides

(12,772, 12,773/CRFS-UEPB): 2 females, Brazil, Pará State, Parauapebas municipality, N4WS-0016 cave, 06°04′35.5″S, 50°11′37.1″W, 21–30.vii.2018, Brandt Meio Ambiente team coll. Additional

records see S1. _Description._ Total length (head + trunk) of specimens 1.43–1.75 mm (n = 5), holotype 1.58 mm. Head. Ratio antennae: trunk = 1: 0.83–0.98 (n = 4), holotype = 1: 0.83; Ant

III larger than Ant II; Ant segments ratio as I: II: III: IV = 1: 1.36–1.77: 1.65–2.03: 2.84–3.27, holotype = 1: 1.72: 1.99: 3.21. Antennal chaetotaxy (no represented): Ant IV dorsally and

ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about two rod sens sub-apical on longitudinal row, ventrally with one subapical-organ and about three

wrinkly sens on longitudinal row (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one

apical PSP, about three wrinkly sens and three smooth mic on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant

II dorsally and ventrally with several short ciliate mic and mac, dorsally with about four sub-apical finger-shaped sens and two subapical rod sens, ventrally with one apical PSP, and

several wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally

with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 42A) with 12 AN (AN1A–3), six A (A0–5), four M

(M1–4), six S (S1–6), two PS (PS2, PS5), four PA (PA1–5), two PM (PM1, PM3), seven PP (PP1–7), and two PE (PE4, PE6) chaetae; A1 as mes, AN1A–3, A0, A2, S5, PA5 and PM3 as mac; interocular P

mic present. Basomedian and basolateral labial fields with A1–5 smooth, M, ME, E and L1–2 ciliate, R reduced (Fig. 42B). Ventral chaetotaxy with 28 ciliate chaetae and one reduced lateral

spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae B.C. present and a collar row of five chaetae distally (Fig. 42B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications.

Labial papilla E with L.P. finger-shaped and subequal the base of apical appendage. Labial proximal chaetae smooth (AN1–3, P2–3) and subequal in length (Fig. 42B). Maxillary palp with T.A.

smooth and 1.58 × larger than B.C. Thorax dorsal chaetotaxy (Fig. 43A). Th II A, M, P series with two mic (A1–2), one mac (A5), three mic (M1–2, M4) and four mic (P4–6E), P3 complex with six

mac, respectively, AL and MS presents. Th III A, M, P series with three mic (A1–3), two mes (A6–7), two mic (M6–6P), three mes (M6E, M7–7E), and one mic (P6), respectively. Ratio Th II: III

 = 0.82–1.13: 1 (n = 6), holotype = 1.13: 1. Abdomen dorsal chaetotaxy (Fig. 43B, C). Abd I A, M series with one (A5) and six (M2–6E) mic, respectively, MS present. Abd II A, M, P series

with two mic (A6–7), two mac (M3, M5), three mic (P5–7) respectively, EL mic and AS present; A5 and M2 bothriotricha surrounded by four and four fan-shaped chaetae, respectively. Abd III A,

M, P series with one mic (A7), three fan-shaped chaetae (A2–3, A6), two mic (M7I–7), three mac (M3, AM6, PM6) and three mic (P6E–7), one mac (P6) chaetae respectively; A5, M2 and M5

bothriotricha with five, two and two fan-shaped chaetae, respectively, AS sens elongated, MS present. Abd IV A–FE series with four mic (A1, A5–6, AE1), one mac (A3), one mic (B1), one mes

(B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, DE3), one mes (D3P), one mic (E4P2), one mes (E4P), three mac (E1–3), one mic (EE12), three mes

(EE9–11), one mic (F1), three mes (F2–3P), one mic (FE2), three mes (FE3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae,

respectively; PS and AS present, and at least six supernumerary sens with uncertain homology _‘s’_ (Fig. 8A); Abd. IV posteriorly with three PSP. Abd V A, M, P series with three mic (A1,

A3), one mes (A6), one mac (A5), two mic (M5A–5E), three mac (M2–3, M5), five mic (P3A–6AE), one mic (P6E), two mes (AP6, PP6), four mac (P1, P3–5) chaetae, respectively; AS and ACC.P4–5

present. Ratio Abd III: IV = 1: 2.98–4.82 (n = 6), holotype = 1: 3.81. Legs. Trochanteral organ diamond shape with about 19 spine-like chaetae, plus 2–3 PSP one external, one on distal

vertex and another (present or absent) on top of posterior spines row of Omt (Fig. 44A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella

wide with two teeth, basal pair subequal, M.T. and A.T. absent. Unguiculus with all lamellae smooth and lanceolate (A.I., A.E., P.I., P.E.) (Fig. 44B); ratio unguis: unguiculus = 1.49–1.80:

1 (n = 6), holotype = 1.80: 1. Tibiotarsal smooth chaetae about 1.25× smaller than unguiculus; tenent hair slightly capitate and about 0.54× smaller than unguis outer lamella. Collophore

(Fig. 44C). Anterior side with 11 ciliate, apically acuminate chaetae, six proximal (thinner); two subdistal and three distal mac; lateral flap with 11 chaetae, five ciliate in the proximal

row and six smooth in the distal row. Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three PSP (Fig. 

44D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 31–39 external and 18–21 internal, external spines larger and thinner than internal ones. Mucro with

three teeth (Fig. 44E), ratio width: length = 0,28 (holotype). _Etymology._ Honor to Dandara Kettley, Brazilian man, transvestite, murdered in 2017 (Ceará, Brazil) in a homophobia crime.

_Remarks. Trogolaphysa dandarae_ SP. NOV. resembles _T. ernesti_, _T. formosensis_ and _T. piracurucaensis_ by chaetae head S5 mac (all other Brazilian cave species with S5 mic); head PM3

mac as in _T. gisbertae_ SP. NOV., but they are different in terms of head ventral proximal collar mac, unguiculus, tenent hair and collophore anterior distal chaetae (5 + 5, smooth PE,

capitate, 3 + 3 and 4 + 4, serrate PE, acuminate, 2 + 2, respectively); Th II P3 complex with 6 + 6 and Th III with 3 + 3 mac (6 + 6 and 0 + 0 in _T. lacerta_ SP. NOV., _T. piracurucaensis_,

_T. ernesti_ and _T. caripensis_); _T. dandarae_ SP. NOV., _T. belizeana_ and _T. jacobyi_ are the only cave species with 3 + 3 teeth in the mucro. See the comparison among them in remarks

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(2019). Article  Google Scholar  Download references ACKNOWLEDGEMENTS R. Zampaulo, R. Andrade, E. Castro, L.M.S.M. Dornellas and M.P.A. Oliveira provided information about specimen’s

collection, cave localities and lithology characterization. Speleological team of Carste, Spelayon, Bioespeleo, Ativo Ambiental, and R. Bessi, R. Andrade, R. Zampaulo, and L.M. Rabelo,

collected the specimens from caves and MSS. FUNDING Senior author is granted by CNPq # 309030/2018-8, J.V.L.C. Oliveira is granted CAPES# 88882.440374/2019-01. B.C.H. Lopes is granted by

CAPES# 88882.440392/2019-01. This research was partially funded by Vallourec Tubos do Brasil Ltda. and Samarco Mineração S.A. AUTHOR INFORMATION AUTHORS AND AFFILIATIONS * Laboratório de

Sistemática de Collembola e Conservação – Coleção de Referência de Fauna de Solo – CCBSA – Universidade Estadual da Paraíba, Campus V, João Pessoa, PB, 58070-450, Brazil Douglas Zeppelini, 

João Victor L. C. Oliveira, Estevam C. Araujo de Lima, Roniere A. Brito, Aila S. Ferreira, Luis C. Stievano, Nathan P. Brito, Misael A. Oliveira-Neto & Bruna C. H. Lopes * Programa de

Pós-graduação em Ciências Biológicas – Zoologia, Universidade Federal da Paraíba, João Pessoa, PB, Brazil Douglas Zeppelini, João Victor L. C. Oliveira & Bruna C. H. Lopes Authors *

Douglas Zeppelini View author publications You can also search for this author inPubMed Google Scholar * João Victor L. C. Oliveira View author publications You can also search for this

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inPubMed Google Scholar CONTRIBUTIONS Senior author contributed to the conception and design of the study, writing the main text, and revision of the whole manuscript. J.V.L.C.O., E.C.A.L.,

R.A.B., and A.S.F. described and illustrated the new species. M.A.O.N. edited the figures and ordered the additional records. L.C.S., N.P.B. performed the slide preparation and ordering the

biological material under analysis. B.C.H.L. organized the biological material and data base. All authors read and approved the final manuscript. CORRESPONDING AUTHOR Correspondence to

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Zeppelini, D., Oliveira, J.V.L.C., de Lima, E.C.A. _et al._ Hotspot in ferruginous rock may have serious implications in Brazilian conservation policy. _Sci Rep_ 12, 14871 (2022).

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