With Special Emphasis on the Equatorial Oceanic Islands

Article Synthesis of the Brazilian Poduromorpha (Collembola: Hexapoda) with Special Emphasis on the Equatorial Oceanic Islands

Estevam C. A. de Lima 1,2,* , Maria Cleide de Mendonça 1, Gabriel Costa Queiroz 1 , Tatiana Cristina da Silveira 1 and Douglas Zeppelini 2

1 Laboratório de Apterygotologia, Departamento de Entomologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro 20940-040, Brazil; [email protected] (M.C.d.M.); [email protected] (G.C.Q.); [email protected] (T.C.d.S.) 2 Laboratório de Sistemática de Collembola e Conservação—Coleção de Referência de Fauna de Solo—CCBSA—Universidade Estadual da Paraíba Campus V, João Pessoa 58070-450, Brazil; [email protected] * Correspondence: [email protected]

Simple Summary: Endemic Collembola species are bioindicators of environmental quality since native species abundance is particularly sensitive to environmental disturbances. Oceanic island biota generally present high percentages of endemic species, and the vulnerability of these species is higher than those of the continents. The objective of this work was to carry out a survey of the Collembola species of the order Poduromorpha in the Brazilian oceanic islands and synthesize a distribution list of this order for Brazil. Our results reveal four new species of Collembola Poduromorpha for

Brazilian oceanic islands that may be useful for the conservation strategies of these island regions and a contributor to the knowledge of the order in Brazil.

Citation: de Lima, E.C.A.; de Mendonça, M.C.; Queiroz, G.C.; da Silveira, T.C.; Abstract: We present new species and records of Poduromorpha for the Brazilian oceanic islands and Zeppelini, D. Synthesis of the Brazilian synthesis of this order in Brazil. Friesea noronhaensis sp. nov., Friesea rochedoensis sp. nov., Willemia Poduromorpha (Collembola: Hexapoda) insularum sp. nov. and Paraxenylla zeliae sp. nov. are described and a diagnosis of the morphospecies with Special Emphasis on the Equatorial Acherontiella sp. Lima and Zeppelini 2015 is provided. We present comparative tables, distribution Oceanic Islands. Insects 2021, 12, 268. and taxonomic keys of the Friesea, Arlesia, Brachystomella, Acherontiella, Paraxenylla, Xenylla, and https://doi.org/10.3390/insects12030268 Willemia found in the Brazilian oceanic islands and their respective congeners recorded in Brazil.

Academic Editor: Steven Trewick Keywords: insular fauna; new species; distribution; Collembola

Received: 26 January 2021 Accepted: 8 March 2021 Published: 22 March 2021 1. Introduction

Publisher’s Note: MDPI stays neutral The order Poduromorpha Börner, 1913 is represented by Collembola with an elongated with regard to jurisdictional claims in body, thorax with three separate segments and prothorax with chaetae. Members of this published maps and institutional afﬁl- order, about 3400 species, occur in all zoogeographic regions [1]. iations. Abrantes et al. [2] summarized a list of Brazilian Collembola with 102 species of Poduromorpha and 2 morphospecies (Mesogastrura cf. ojcoviensis and Mesaphorura sp. gr. atlantica), none of them recorded in Brazilian oceanic islands. Oceanic islands generally have a high percentage of endemic species. However, in Brazil, the only survey of the Collembola fauna in oceanic islands was done by Lima Copyright: © 2021 by the authors. Licensee MDPI, Basel, Switzerland. and Zeppelini in 2015 [3], who presented 36 new records of Collembola, among them This article is an open access article 4 species and 3 morphospecies of the Poduromorpha order: Friesea cubensis Potapov and distributed under the terms and Banasko, 1985; Brachystomella agrosa Wray, 1953; Xenylla yucatana Mills, 1938; Isotogastrura conditions of the Creative Commons mucrospatulata Palacios-Vargas, Lima and Zeppelini, 2013; Acherontiella sp.; Arlesia sp. and Attribution (CC BY) license (https:// Willemia sp., all from Fernando de Noronha Archipelago. creativecommons.org/licenses/by/ We present results of recent surveys of the Archipelago of São Pedro e São Paulo, 4.0/). Atol das Rocas (Rocas Atoll) and the taxonomic advances for Fernando de Noronha

Insects 2021, 12, 268. https://doi.org/10.3390/insects12030268 https://www.mdpi.com/journal/insects Insects 2021, 12, 268 2 of 39

Archipelago after Lima and Zeppelini [3]. Friesea noronhaensis sp. nov., Friesea rochedoensis sp. nov., Willemia insularum sp. nov. and Paraxenylla zeliae sp. nov are described. We provide comparative tables, distribution, remarks and taxonomic keys to Brazilian species of the genera Friesea, Arlesia, Brachystomella, Acherontiela, Paraxenylla, Xenylla, Willemia, Isotogastrura, as well as an update of the distribution and habitat synthesis of the Brazilian Poduromorpha, including contributions after Abrantes et al. [2].

2. Materials and Methods 2.1. Poduromorpha Survey of the Brazilian Oceanic Islands This survey was conducted in 3 Brazilian oceanic islands (2 Archipelagos and 1 Atoll) located in the Equatorial Atlantic Ocean, all belonging to the Brazilian national territory, as follows: Fernando de Noronha Archipelago—AFN (3◦500 S, 32◦150 O), São Pedro e São Paulo Archipelago—ASPSP (0◦550 N, 29◦200 O) and Atol das Rocas (Rocas Atoll)—AT (3◦510 S, 33◦480 O) about 360, 1010 and 267 km from the American continent, respectively (Figure1).

2.2. Sample Procedure The specimens were collected in three coastal environmental areas: the intertidal zone in sand or rock areas, without vegetation, referred to as sandy beach (SB); the sloping land closest to the intertidal zone, with vegetation, referred to as slope forest (SF) and the forest more distant from the beach, referred to as top forest (TF), this latest area only exists in Fernando de Noronha. Specimens were preserved in ethanol, cleared in Nesbitt’s ﬂuid and then mounted on glass slides in Hoyer’s medium [4]. Figures were elaborated in computer graphic software, photographs and measurements were obtained using a Zeiss Axio Scope A1 microscope with Axiocam 105 color, and Zen 2 Blue software, and deposited in the Coleção de Re- ferência de Fauna de Solo at the Universidade Estadual da Paraíba, João Pessoa, PB, Brazil.

2.3. Taxonomic Procedure The nomenclature of furcal development stages follows Cassagnau 1958. Antennal S-chaetae are numbered from S1 to S8 [5]. Dorsal chaetotaxy nomenclature follows the chaetal group system [6–8].

2.4. Species List of Brazilian Poduromorpha Species records, collection location, habitat and biotope information for each species were modified from [2], included: newly described species, new records published between 2012 and 2019 and a survey of the Brazilian oceanic islands between 2012 and 2017. Publications that do not provide identifications to continental species were omitted except genus and morpho-species from Brazilian oceanic islands. Information on the world distribution of species was based on [9] and original new records. Biogeographical distribution regions according to [10], modified by [9,11] as follows: Boreal (Bor) include regions 1–8, Neotropical (Neo) regions 24–30, South African (SAf) region 31, Paleotropical (Pal) regions 9–23, Australian (Aus) regions 32–34, and Antarctic (Ant) regions 35–37. Species restricted to Brazil were based on Culik and Zeppelini [9], defining the Brazilian biogeographic sub- regions: Amazon (Amz), North and Central Brazil (NCB) and Pampa (Pam) corresponding to biogeographic regions 26, 27 and 29, respectively [9]. Species distributed, at least, in four of the major regions (Neo, Pal, etc.) are considered cosmopolitan (Cos).

2.5. Abbreviations Abd—abdominal segment, Ant—antennal segment, Cd—cephalic diagonal, PAO— postantennal organ, Oc—ocular, ta—anterior trichobothria, Th—thoracic segment, Tita— tibiotarsus, tm—medial trichobothria, tl—lateral trichobothria, tp—posterior trichobothria, Di—dorsointernal group of chaetae, De—dorsoexternal group of chaetae, DL—dorsolateral group of chaetae, CRFS-UEPB—Coleção de Referência da Fauna de Solo-Universidade Insects 2021, 12, x FOR PEER REVIEW 3 of 40

Insects 2021, 12, 268 3 of 39

corresponding to biogeographic regions 26, 27 and 29, respectively [9]. Species distributed, at least, in fourEstadual of the da major Paraíba, regions IUCN—International (Neo, Pal, etc.) Union are for considered Conservation cosmopolitan of Nature. Brazilian (Cos). states abbreviations according to Table1.

FigureFigure 1. Brazilian1. Brazilian oceanic oceanic island ofisland the Equatorial of the Equatorial Atlantic Ocean. Atlant Mapic show Ocean. distribution Map show of sites distribution sampled in Fernando of sites de Noronhasampled archipelago, in Fernando Atol dasde RocasNoronha (Rocas archipelago, Atoll) and Sã oAtol Pedro das e S ãRocaso Paulo (Rocas Archipelago. Atoll) and São Pedro e São Paulo Archipelago.

2.5. Abbreviations Abd––abdominal segment, Ant––antennal segment, Cd––cephalic diagonal, PAO–– postantennal organ, Oc––ocular, ta––anterior trichobothria, Th––thoracic segment, Tita–– tibiotarsus, tm––medial trichobothria, tl––lateral trichobothria, tp––posterior trichoboth-

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ria, Di––dorsointernal group of chaetae, De––dorsoexternal group of chaetae, DL––dorsolateral group of chaetae, CRFS-UEPB––Coleção de Referência da Fauna de Solo-Univer- sidade Estadual da Paraíba, IUCN––International Union for Conservation of Nature. Bra- Insects 2021, 12, 268 zilian states abbreviations according to Table 1. 4 of 39

Table 1. Abbreviations of names of the Brazilian states and oceanic islands. Number of Poduromopha species recorded and region location of each state and oceanic island. Brazilian continental region designations: N––north, NE––northeast, CWTable––west 1. Abbreviations (central part), SE of–– namessoutheast, of the S Brazilian––south. Brazilia states andn insular oceanic region: islands. EA–– Numberequatorial of Poduromopha Atlantic; SA––South species Atlantic. recorded Mapand shows region the location geographic of each position state and of Brazilian oceanic island. states and Brazilian oceanic continental islands. region designations: N—north, NE—northeast, CW—west (central part), SE—southeast, S—south. Brazilian insular region: EA—equatorial Atlantic; SA—South Atlantic. Abbreviation––State Species Region Abbreviation––Oceanic Island Species Region Map shows the geographic position of Brazilian states and oceanic islands. AC––Acre 0 N FN––Fernando de Noronha 7 EA ALAbbreviation—State––Alagoas Species0 RegionNE Abbreviation—OceanicAR––Atol das Rocas Island Species2 RegionEA AM––AmazonasAC—Acre 24 0N SPSP FN—Fernando––São Pedro de Noronhae São Paulo 71 EA AP––AL—AlagoasAmapá 2 0 NEN TR–– AR—AtolTrindade das e RocasMartim Vaz 20 EASA BAAM—Amazonas––Bahia 1 24NE N SPSP—São Pedro e São Paulo 1 EA CEAP—Amap––Ceará á 1 2NE N TR—Trindade e Martim Vaz 0 SA ES––EspiroBA—Bahia Santo 17 1 NESE CE—Ceará 1 NE GO––Goiás 0 CW ES—Espiro Santo 17 SE MA––GO—GoiMaranhãoás 0 0 CWNE MG––MA—MaranhMinas Geraisão 17 0 NESE MS––MatoMG—Minas Grosso Geraisdo Sul 1 17CW SE MTMS—Mato––Mato GrossoGrosso do Sul5 1CW CW MT—Mato Grosso 5 CW PA––PA—ParPará á 11 11N PB––PB—ParaParaíbaí ba4 4NE PE––PE—PernambucoPernambuco 6 6NE PI––PI—PiauPiauí í 2 2 NE PR—Paraná 3 SE PRRJ—Rio––Paraná de Janeiro 3 80SE RJRN—Rio––Rio de Grande Janeiro do Norte80 5 NESE RN––Rio RO—RondGrande doônia Norte 5 1NE N RO––RR—RoraimaRondônia 1 0N RS—Rio Grande do Sul 0 S RR––Roraima 0 N SC—Santa Catarina 0 S RS––Rio GrandeSE—Sergipe do Sul 0 4 NES SC––SantaSP—S Catarinaão Paulo 0 9 SES SETO—Tocantins––Sergipe 4 0NE N SP––São Paulo 9 SE TO––Tocantins 3. Results0 N The Brazilian Poduromorpha fauna is recorded in the following states and oceanic 3. islandsResults (Table 1). The Brazilian Poduromorpha fauna is recorded in the following states and oceanic 3.1. Poduromorpha Survey of the Brazilian Equatorial Oceanic Islands islands (Table 1). The Poduromorpha fauna of the Brazilian oceanic islands is represented by 9 species, 3.1.as Poduromorpha follows: Survey of the Brazilian Equatorial Oceanic Islands

3.1.1.The Neanuridae Poduromorpha Börner, fauna 1901 of the Brazilian oceanic islands is represented by 9 species, as follows: Friesea noronhaensis sp. nov. Lima and Zeppelini 3.1.1. NeanuridaeType material. Börner,Holotype 1901male on the slide, Brazil, Fernando de Noronha Archipelago, ◦ 0 00 ◦ 0 00 FrieseaRata noronhaensis Island, (3 48 sp.45.61 nov.S; Lima 32 23 and26.17 ZeppeliniO), SF, 19.vii.2012, E.C.A. Lima and A.S. Ferreira leg., deposited at the CRFS-UEPB # 14574. Paratypes: 2 females and 2 males on slides, Type material. Holotype male on the slide, Brazil, Fernando de Noronha Archipelago, same data as holotype, deposited at the CRFS-UEPB # 14572, 14573, 15060, 1563. Rata Island, (3°48′45.61″ S; 32°23′26.17″ O), SF, 19.vii.2012, E.C.A. Lima and A.S. Ferreira Description. Habitus typical of the genus. Color in ethanol dark gray. Body with leg., deposited at the CRFS-UEPB # 14574. Paratypes: 2 females and 2 males on slides, secondary integuments. Body length average: 512 µm (n = 5); holotype measurements in same data as holotype, deposited at the CRFS-UEPB # 14572, 14573, 15060, 1563. Table2. Antenna. Antenna shorter than cephalic diagonal (Table2). Ant IV with 6 S-chaetae (S1–4, S7–8), 1 small ventrolateral S-microchaeta and 1 small subapical organite. Apical bulb simple; with 5 dorsal clavate chaetae and about 12 smooth chaetae ventrally. Sensory organ of Ant III consisting of 2 small subcylindrical, bent, internal S-chaetae covered by a fold of the integument, 2 subcylindrical guard S-chaetae Sgd and Sgv, and 1 ventral S-microchaeta. Ant II with 12 chaetae; Ant I with 7 chaetae (Figure2a,b). Insects 2021, 12, x FOR PEER REVIEW 5 of 40

Description. Habitus typical of the genus. Color in ethanol dark gray. Body with secondary integuments. Body length average: 512 µm (n = 5); holotype measurements in Table 2.

Table 2. Holotype measurements for Friesea noronhaensis sp. nov., Friesea rochedoensis sp. nov., Wil- lemia insularum sp. nov., Acherontiella sp. and Paraxenylla zeliae sp. nov. All measurements in µm.

Character F. noronhaensis F. rochedoensis W. insularum Acherontiella. sp. P. zeliae Body 528.57 1106.69 520.64 746.69 710.50 Cd 135.52 234.98 78.58 130.36 149.12 Ant I 14.82 33.42 8.93 12.31 14.51 Ant II 16.23 30.08 14.18 16.47 18.75 Ant III 11.97 19.75 16.38 16.38 26,85 Ant IV 19.03 23.07 11.90 31.92 32.63 Mucro 4.52 6.70 Absent Absent 27.41 Dens 10.41 23.49 Absent Absent 28.02 Manubrium 22.96 23.87 Absent Absent 36.28 Unguis III 12.50 20.28 10.57 13.35 18.27 Insects 2021, 12, 268 5 of 39 Tita III 22.26 34.61 10.91 18.55 28.40 Femur III 19.82 32.36 17.73 18.09 15.82

Table 2. HolotypeTr measurements for15.55Friesea noronhaensis sp.24.61 nov., Friesea rochedoensis9.08 sp. nov., Willemia15.50 insularum sp. nov.,10.32 Acherontiella sp.Cx and Paraxenylla zeliae32.53sp. nov. All measurements36.09 in µm. 12.63 23.92 17.34 Th II ordinary Character F. noronhaensis4.98 F. rochedoensis13.16 W. insularum4.19 Acherontiella.7.89 sp. P. zeliae8.02 chaetae Body 528.57 1106.69 520.64 746.69 710.50 ThCd II s-chaetae 135.5211.83 234.9816.17 78.587.98 130.3613.53 149.1214.22 AntAnal I spine 14.8212.78 33.4220.66 8.936.97 A 12.31bsent Absent 14.51 AbdAnt V II chaetae a1 16.23 30.08 14.18 16.47 18.75 Ant III 11.975.25 19.7519.56 16.389.65 16.388.02 26,858.21 Ant IVor ord. 19.03 23.07 11.90 31.92 32.63 AbdMucro V chaetae p1 4.52 6.70 Absent Absent 27.41 13.09 23.10 9.98 8.13 15.03 Densor Mac. 10.41 23.49 Absent Absent 28.02 Manubrium 22.96 23.87 Absent Absent 36.28 UnguisAbd V III s-chaetae 12.5015.08 20.2820.35 10.5721.70 13.3517.08 18.2716.31 Tita III 22.26 34.61 10.91 18.55 28.40 Femur III 19.82 32.36 17.73 18.09 15.82 TrAntenna. Antenna 15.55 shorter than 24.61cephalic diagonal 9.08 (Table 2). Ant 15.50 IV with 6 S 10.32-chaetae (S1–Cx4, S7–8), 1 small ventrolateral 32.53 S- 36.09microchaeta and 12.63 1 small subapical 23.92 organite. 17.34 Apical Th II ordinary chaetae 4.98 13.16 4.19 7.89 8.02 Thbulb II s-chaetae simple; with 5 dorsal 11.83 clavate chaetae 16.17 and about 12 7.98 smooth chaetae 13.53 ventrally. 14.22Sensory organAnal spine of Ant III consisting 12.78 of 2 small 20.66subcylindrical, 6.97bent, internal S Absent-chaetae covered Absent by a Abd V chaetae a1 or ord. 5.25 19.56 9.65 8.02 8.21 Abd Vfold chaetae of p1 the or Mac. integument, 13.09 2 subcylindrical 23.10 guard S-chaetae 9.98 Sgd and Sgv, 8.13 and 1 ventral 15.03 S- Abdmicrochaeta. V s-chaetae Ant II with 15.08 12 chaetae; Ant 20.35 I with 7 chaetae 21.70 (Figure 2a,b 17.08). 16.31

Figure 2. Friesea noronhaensisFigure 2. Friesea sp. noronhaensisnov. (a) dorsalsp. nov. view (a) dorsal of antenna view of antenna (b) ventral (b) ventral view view of antenna of antenna.. Scale Scale bar: 10 µm. bar: 10 µm. Head. Eyes 8 + 8 in dark ocular plate; PAO absent; chaetae a0, d0 and 3 Oc present (Figure3). Pre-labral/labral formula: 4/3,3,4. Mandible with seven teeth, three apical, a medial and three basal (Figure4a). Maxilla typical of the genus (Figure4b). Labium with papillated chaeta L (Figure4c). Head and body chaetotaxy composed of slightly rugged acuminate ordinary chaetae. S-chaetae smooth, except on Th II–III in DL that are roughly blunt. S-chaetae measuring about twice the size of anterior ordinary chaetae in Th II–III and Abd I–IV. Abd V–VI with posterior strong rough, blunt chaetae longer than others of the body. Sensillar formula by half tergum: 022/11111 (Figures3 and5). Thoracic chaetotaxy. Th I: 4 + 4; Th II: 13 + 13 (Di: 4 chaetae; De: 4 chaetae + 1 S-chaetae; Dl: 2 chateae, 1 S-chaetae +1 ms); Th III: 11 + 11 (Di: 3 chaetae; De: 4 chaetae +1 S-chaetae; Dl: 2 chaetae +1 S-chaetae) see Figure3. Insects 2021, 12, x FOR PEER REVIEW 6 of 40

Head. Eyes 8 + 8 in dark ocular plate; PAO absent; chaetae a0, d0 and 3 Oc present Insects 2021, 12, 268 (Figure 3). Pre-labral/labral formula: 4/3,3,4. Mandible with seven teeth, three apical,6 of a39 medial and three basal (Figure 4a). Maxilla typical of the genus (Figure 4b). Labium with papillated chaeta L (Figure 4c).

Figure 3 3.. FrieseaFriesea noronhaensis noronhaensis sp.sp. nov nov,, d dorsalorsal view view of of head head and and Th Th I– I–III.III. Scale Scale bar bar 10 10 µm.µm.

Abdominal chaetotaxy. Abd I–III: 12 + 12 (Di: 3; De: 4 + 1 S-chaetae; Dl: 4). Abd IV: 11 + 11 (Di: 4; De: 2 + 1 S-chaetae; Dl: 4). Abd V: 6 + 6 (3 anterior acuminate chaetae and 3 posterior blunt chaetae measuring about twice the size than anterior plus 1 + 1 S-chaetae Insects 2021, 12, 268 7 of 39

in p2) chaetae. Abd VI with 8 clavate chaetae and 3 anal spines (Figure5), 2 in a1 position and the third as MO. Appendages. Tita I–III: 18, 18, 17 chateae, with 3, 4, 4 longer and clavate tenent hair, respectively (Figure6a); Femora I–III: 10, 14, 14 chaetae; trochanters I–III: 5, 5, 5 chaetae; coxae I–III: 3, 7, 7 chaetae; subcoxae I–III with 0, 2, 2 chaetae; epicoxae I–III with 1, 2, 2. Unguis without teeth. Ventral tube with 4 + 4 chaetae. Tenaculum with 2 + 2 teeth and furca Insects 2021, 12, x FOR PEER REVIEWstate 1 (Figure6b). Genital plate of female and male with 9 and 28 chaetae, respectively. 7 of 40

Anal valves with 16 + 16 chaetae, each anal valve with 2 hr chaetae (Figure6c).

FigureFigure 4. 4.Friesea Friesea noronhaensis noronhaensissp. sp. nov. nov (.a ()a mandible,) mandible scale, scale bar: bar: 5 µ5 m.µm. (b ()b maxilla,) maxilla scale, scale bar: bar: 5 µm. (c) 5labiumµm. (c), scale labium, bar: scale 10 µm. bar: 10 µm.

EtymologyHead. Species and body name cha derivesetotaxy fromcomposed the type of slightly locality, rugged popularly acuminate known asordinary Noronha. chaetae. Remarks.S-chaetaeFriesea smooth, noronhaensis except on sp.Th II nov.–III isinsimilar DL that to areF. cubensisroughlyPotapov blunt. S- andchaetae Banasco, measuring 1985; F.about furculata twiceDeharveng the size of anterior and Bedos, ordinary 1991; chF.a rubenietae in DeharvengTh II–III and and Abd Bedos I–IV. 1991Abd V and–VI with F. australicaposteriorGreenslade strong rough and, Deharveng,blunt chaetae 1997 longer by developedthan others furca,of the densbody. with Sensillar 3 chaetae formula by and separatehalf tergum: mucro. 022/11111Friesea (Fig cubensisures and3 and the 5).new species presents 3 anal spines, while Thoracic chaetotaxy. Th I: 4 + 4; Th II: 13 + 13 (Di: 4 chaetae; De: 4 chaetae + 1 S- chaetae; Dl: 2 chateae, 1 S-chaetae +1 ms); Th III: 11 + 11 (Di: 3 chaetae; De: 4 chaetae +1 S- chaetae; Dl: 2 chaetae +1 S-chaetae) see Figure 3. Abdominal chaetotaxy. Abd I–III: 12 + 12 (Di: 3; De: 4 + 1 S-chaetae; Dl: 4). Abd IV: 11 + 11 (Di: 4; De: 2 + 1 S-chaetae; Dl: 4). Abd V: 6 + 6 (3 anterior acuminate chaetae and 3 posterior blunt chaetae measuring about twice the size than anterior plus 1 + 1 S-chaetae in p2) chaetae. Abd VI with 8 clavate chaetae and 3 anal spines (Figure 5), 2 in a1 position and the third as MO.

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Insects 2021, 12, 268 8 of 39 Appendages. Tita I–III: 18, 18, 17 chateae, with 3, 4, 4 longer and clavate tenent hair, respectively (Figure 6a); Femora I–III: 10, 14, 14 chaetae; trochanters I–III: 5, 5, 5 chaetae; coxae I–III: 3, 7, 7 chaetae; subcoxae I–III with 0, 2, 2 chaetae; epicoxae I–III with 1, 2, 2. F. furculata, F. rubeni and F. australica feature 0, 2 and 4, respectively. The new species differs Unguis without teeth. Ventral tube with 4 + 4 chaetae. Tenaculum with 2 + 2 teeth and from F. cubensis by posterior chaetae blunt versus acuminate in Abd V–VI and tenent hair furcaclavate state 1 versus (Figure acuminate. 6b). Genital For morphologicalplate of female comparisons and male with between 9 andFriesea 28 chaetae,species respec- recorded tively.in Brazil,Anal valves see Table with3. 16 + 16 chaetae, each anal valve with 2 hr chaetae (Figure 6c).

FigureFigure 5. Friesea 5. Friesea noronhaensis noronhaensis sp. nov.sp. nov. 8, dorsal 8, dorsal view view of abdominal of abdominal segments segments I–VI. I–VI. Scale Scale bar bar 10 10µm.µm.

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Figure 6. Friesea noronhaensis sp. nov. (a) Tita III, scale bar: 10 µm. (b) furca, scale bar: 10 µm. (c) Figure 6. Friesea noronhaensis sp. nov. (a) Tita III, scale bar: 10 µm. (b) furca, scale bar: 10 µm. ventral view(c )of ventral abdominal view of abdominalsegment segmentII–VI, scale II–VI, bar: scale 20 bar: µm. 20 µ m.

EtymologyDistribution. Species .nameFriesea noronhaensisderives fromsp. nov. the is type registered locality, only in popularly the Fernando known de Noronha as Noro- nha. Archipelago on Fernando de Noronha Island (SF sites) and Rata Island (SF and TF sites). Remarks.The specimens Friesea noronhaensis were collected sp. in superﬁcial nov. is similar soil and leaf.to F. cubensis Potapov and Banasco, 1985; F. furculata Deharveng and Bedos, 1991; F. rubeni Deharveng and Bedos 1991 and F. australica Greenslade and Deharveng, 1997 by developed furca, dens with 3 chaetae and separate mucro. Friesea cubensis and the new species presents 3 anal spines, while F. furculata, F. rubeni and F. australica feature 0, 2 and 4, respectively. The new species differs from F. cubensis by posterior chaetae blunt versus acuminate in Abd V–VI and tenent hair clavate versus acuminate. For morphological comparisons between Friesea species recorded in Brazil, see Table 3. Distribution. Friesea noronhaensis sp. nov. is registered only in the Fernando de Noro- nha Archipelago on Fernando de Noronha Island (SF sites) and Rata Island (SF and TF sites). The specimens were collected in superficial soil and leaf.

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Table 3. Morphological characters of Friesea species record from Brazil. S Abd VI—number of the spine on Abd VI; M Abd VI—modiﬁed chaetae on Abd VI; Ten h—tenent hair; BD—body length in µm; AP—apical vesicle; TL—type locality; “?”—missing date; number in Furca column refers of furca “State” according to [12].

Species Eyes S Abd VI Furca M Abd VI Ten h BD AP Th I Tita I-III TL Color F. arlei Massoud and Bellinger, 1963 4 + 4 Absent 2 Clavate 1 clavate ? simple 2 + 2 ? Jamaica Blue F. boitata Queiroz and Mendonça, 2015 8 + 8 Absent 5 Clavate 4,5,5 clavete 400–800 simple 4 + 4 18,18,17 Brazil Blue F. claviseta Axelson, 1900 8 + 8 3 2 Clavate ? 550–700 Finland ? F. cubensis Potapov and Banasko, 1985 8 + 8 3 1 Acuminate Acuminate 670–1000 simple 4 + 4 ? Cuba ? F. curupira Queiroz and Mendonça, 2015 1 + 1 2 5 Acuminate Acuminate 450–470 simple 4 + 4 15,15,14 Brazil Dark blue F. josei Palacios-Vargas, 1986 8 + 8 Absent 5 Spinne 3,3,3 clavate 1100 simple 3 + 3 17,17,16 Porto Rico Blue F. jurubatiba Silveira and Mendonça, 2018 8 + 8 Absent 5 Clavate 4,5,5 clavate 540–680 simple 4 + 4 18,18,17 Brazil Dark gray F. magnicornis Denis, 1931 8 + 8 3 * 2 Clavate 5 clavate 900 ? ? ? Costa Rica Gray F. mirabilis (Tullberg, 1871) 8 + 8 3 2 Acuminate ? ? ? ? ? ? ? F. noronhaensis sp. nov. 8 + 8 3 1 Clavate 3,4,4 clavate 440–530 simple 4 + 4 18,18,17 Brazil Dark blue F. reducta Denis, 1931 8 + 8 Absent 5 Clavate 4,5,5 clavete 600 simple 3 + 3 ? Costa Rica Gray F. rochedoensis sp. nov. 8 + 8 3 1 Acuminate 3,4,4 clavate 890–1180 simple 4 + 4 18,18,17 Brazil Light blue F. sublimis Macnamara, 1921 8 + 8 3 2 Clavate ? ? ? ? ? ? ? * Different sizes. Insects 2021, 12, x FOR PEER REVIEW 11 of 40

Friesea rochedoensis sp. nov. Lima and Zeppelini Type material. Holotype female on slide, Brazil, São Pedro e São Paulo Archipelago, Insects 2021, 12, 268 11 of 39 Belmonte Island, (0°54′59.98″ N; 29°20′44.03″ O), guano over rocks, May 2017, E.C.A. Lima and D. Zeppelini leg., deposited at the CRFS-UEPB # 14571. Paratypes: 2 males on slides, Brazil, São Pedro e SãoFriesea Paulo rochedoensis Archipelago,sp. nov. Lima Belmonte and Zeppelini Island, (0°54′59.74″ N; 29°20′44.57″ O), guano over rocks, 24 AprilType material 2017,. Holotype E.C.A. femaleLima onand slide, D. Brazil,Zeppelini São Pedro leg., e Sdepositedão Paulo Archipelago, at the Belmonte Island, (0◦54059.9800 N; 29◦20044.0300 O), guano over rocks, May 2017, E.C.A. CRFS-UEPB # 14569, 14570.Lima and D. Zeppelini leg., deposited at the CRFS-UEPB # 14571. Paratypes: 2 males Description. Habituson slides, typical Brazil, of the São genus. Pedro e SLightão Paulo blue Archipelago, color in Belmonteethanol. Island,Body (0with◦54059.74 well00 -N; developed secondary 29integuments.◦20044.5700 O), guano Body over length rocks, average: 24 April 2017,995 µm E.C.A. (n Lima = 5); and holotype D. Zeppelini meas- leg., deposited at the CRFS-UEPB # 14569, 14570. urements as in Table 2. Description. Habitus typical of the genus. Light blue color in ethanol. Body with Antenna. Antennawell-developed shorter than secondary cephalic integuments. diagonal Body (Table length 2). average:Ant IV 995dorsallyµm (n = with 5); holotype 6 S- chaetae (S1–4, S7–8), 1measurements small ventrolateral as in Table2 .S-microchaeta and 1 small subapical organite; Antenna. Antenna shorter than cephalic diagonal (Table2). Ant IV dorsally with apical bulb simple ventrally;6 S-chaetae with (S1–4, 3 S7–8),–4 dorsoapical 1 small ventrolateral and 10 S-microchaeta–12 clavate and plus 1 small 5 smooth subapical ven- organ- troapical chaetae. Sensoryite; apical organ bulb of simple Ant ventrally; III consisting with 3–4 of dorsoapical 2 small andsubcylindrical, 10–12 clavate plus bent, 5 smooth internal S-chaetae coveredventroapical by a fold chaetae. of the Sensory integument, organ of Ant2 subcylindrical III consisting of 2guard small subcylindrical, S-chaetae Sgd bent, and Sgv, and 1 ventralinternal S-microch S-chaetaeaeta. covered (Fig byure a fold7a,b of). the Ant integument, I with 7 2 ch subcylindricalaetae; Ant guard II with S-chaetae 12 Sgd and Sgv, and 1 ventral S-microchaeta. (Figure7a,b). Ant I with 7 chaetae; Ant II with chaetae. 12 chaetae.

FigureFigure 7. 7. FrieseaFriesea rochedoensisrochedoensissp. nov.sp. (nova) dorsal. (a) viewdorsal of antenna, view of scale antenna bar: 20 ,µ scalem. (b) ventralbar: 20 view µm. of ( antenna,b) ventral scale view bar: µ of antenna20 m. , scale bar: 20 µm. Head (Figure8a). Eyes 8 + 8 in dark ocular plate; PAO absent; chaetae a0, d0 and 3 Oc present. Pre-labral/labral formula: 4/3,3,4. Mandible with seven teeth, three apical, a Head (Figure 8a).medial Eyes and 8 + three 8 in basal dark (Figure ocular8b). Maxillaplate; typicalPAO absent; of the genus. cha Labiumetae a0, with d0 papillated and 3 Oc present. Pre-labral/labralchaeta L. formula: 4/3,3,4. Mandible with seven teeth, three apical, a medial and three basal (FigureHead and 8b body). Maxilla chaetotaxy typical composed of the of slightlygenus. rough Labium acuminate with ordinary papillated chaetae and S-chaetae about 1/3 longer than ordinary chaetae in Th I–II. Abd I–V S-chaetae chaeta L. subequal or slightly longer than ordinary posterior chaetae. Sensillar formula by half Head and body chaetotaxytergum: 022/11111. composed of slightly rough acuminate ordinary chaetae and S-chaetae about 1/3 longer than ordinary chaetae in Th I–II. Abd I–V S-chaetae subequal or slightly longer than ordinary posterior chaetae. Sensillar formula by half tergum: 022/11111. Thoracic chaetotaxy (Figure 8a). Th I: 4 + 4; Th II: 13 + 13 (Di: 4 chaetae; De: 4 chaetae + 1 S-chaetae; Dl: 2 chaetae, 1 S-chaetae + 1 ms); Th III: 11 + 11 chaetae (Di: 3 chaetae; De: 4 chaetae +1 S-chaetae; Dl: 2 chaetae +1 S-chaetae). Abdominal chaetotaxy (Figure 8c). Abd I–IV: 12 + 12 (Di 3; De 5; Dl 4, posterior chaetae about 1/4 longer than size anterior chaetae). Abd V: 6 + 6 (3 anterior chaetae and 3 posterior of which one S-chaetae subequal chaetae). Abd VI with 6 chaetae and 3 anal spines.

Insects 2021, 12, 268 12 of 39 Insects 2021, 12, x FOR PEER REVIEW 12 of 40

FigureFigure 8.8. FrieseaFriesea rochedoensis rochedoensis sp.sp. nov nov.. (a) dorsal (a) dorsal view viewof head of and head Th andI–III, Th scale I–III, bar: scale 20 µm. bar: (b)20 µm. (bmandible) mandible,, scale scale bar: bar: 20 µm 20 µ. m.(c) d (corsal) dorsal view view of abdominal of abdominal segments segments I–VI, I–VI, scale scale bar: 20 bar: µm. 20 µm.

Insects 2021, 12, 268 13 of 39

Thoracic chaetotaxy (Figure8a). Th I: 4 + 4; Th II: 13 + 13 (Di: 4 chaetae; De: 4 chaetae + 1 S-chaetae; Dl: 2 chaetae, 1 S-chaetae + 1 ms); Th III: 11 + 11 chaetae (Di: 3 chaetae; De: 4Insects chaetae 2021 +1, 12 S-chaetae;, x FOR PEER Dl: 2REVIEW chaetae +1 S-chaetae). 13 of 40 Abdominal chaetotaxy (Figure8c). Abd I–IV: 12 + 12 (Di 3; De 5; Dl 4, posterior chaetae about 1/4 longer than size anterior chaetae). Abd V: 6 + 6 (3 anterior chaetae and 3 posterior of which one S-chaetae subequal chaetae). Abd VI with 6 chaetae and 3 anal spines. Appendages. Tita I–III: 18, 18, 17 chaetae, with 3, 4, 4 longer and clavate tenent hairs, Appendages. Tita I–III: 18, 18,respectively; 17 chaetae, with Femora 3, 4, 4 longer I–III: and 10, clavate 11, 10 tenentchaetae; hairs, trochanters I–III: 5, 5, 5 chaetae; coxae I–III: respectively; Femora I–III: 10, 11, 103, chaetae; 7, 7 ch trochantersaetae; subcoxae I–III: 5, 5, I 5–III chaetae; with coxae 0, 2, I–III:2 ch 3,aetae; epicoxae I–III: 0, 2, 2. Unguis short, 7, 7 chaetae; subcoxae I–III with 0, 2, 2 chaetae; epicoxae I–III: 0, 2, 2. Unguis short, without tooth on inner edge. Ventral tubewithout with 4 + 4tooth chateae. on inner Tenaculum edge. with Ventral 2 + 2 tube teeth with and 4 + 4 chateae. Tenaculum with 2 + 2 teeth furca type 1. Genital plate of femaleand (Figure furca9a) type and 1. male Genital with 17–19plate andof female26–28 chaetae(Figure 9a) and male with 17–19 and 26–28 chae- (Figure9b). Anal valves with 16 + 16tae and (Fig 2 hrure chaetae 9b). Anal (Figure valves9c). with 16 + 16 and 2 hr chaetae (Figure 9c).

FigureFigure 9. Friesea 9. rochedoensis Friesea rochedoensissp. nov. (a) femalesp. nov. genital (a) female plate, scale genital bar: 20plateµm., (scaleb) male bar: genital 20 µm. plate, (b) male genital plate, scale bar: 20 µm. (c) scale bar:left 20 analµm. valve (c) left, analscale valve, bar: scale20 µm. bar: 20 µm.

Etymology. Species name derives from the old name of the type locality that was previously known as “Rochedo de São PedroEtymology e São Paulo”.. Species name derives from the old name of the type locality that was pre- Remarks. F. rochedoensis sp. nov.viously is similar knownto F. noronhaensis as “Rochedosp. nov. de by São chaetotaxy, Pedro e eyes São Paulo”. number, mandible, furca type, number ofRemarks anal spines. F. and rochedoensis clavate tenent sp. hair.nov. Differs is similar from to F. noronhaensis sp. nov. by chaetotaxy, F. noronhaensis by color (light blue vs.eyes dark number, gray), for mandible, absent posterior furca blunt type, clavate number chaetae of anal spines and clavate tenent hair. Differs in Abd V–VI, by the size (body lengthfrom 995 F. vs.noronhaensis 512 µm) and by a numbercolor (light of clavate blue chaetaevs. dark gray), for absent posterior blunt clavate dorsal and ventroapical of the Ant IV (3–4, 10–12 vs. 5, 0). Friesea rochedoensis sp. nov. chaetae in Abd V–VI, by the size (body length 995 vs. 512 µm) and a number of clavate chaetae dorsal and ventroapical of the Ant IV (3–4, 10–12 vs. 5, 0). Friesea rochedoensis sp. nov. has hardly differentiated S-chaetae (especially on Abd V) from ordinary chaetae, whereas F. noronhaensis sp. nov. and F. cubensis S-chaetae are easily differentiated from ordinary chaetae. Friesea rochedoensis sp. nov. has chaetae a1 and p1 subequal in Abd V, while F. noronhaensis sp. nov and F. cubensis have p1 greater than a1 in Abd V (Figure 10a–c). For others, morphological comparisons between Friesea species recorded in Brazil, see Table 3.

Insects 2021, 12, 268 14 of 39

has hardly differentiated S-chaetae (especially on Abd V) from ordinary chaetae, whereas F. noronhaensis sp. nov. and F. cubensis S-chaetae are easily differentiated from ordinary chaetae. Friesea rochedoensis sp. nov. has chaetae a1 and p1 subequal in Abd V, while F. noronhaensis sp. nov and F. cubensis have p1 greater than a1 in Abd V (Figure 10a–c). For others, morphological comparisons between Friesea species recorded in Brazil, see Table3. Insects 2021, 12, x FOR PEER REVIEW 14 of 40 Insects 2021, 12, x FOR PEER REVIEW Distribution. Friesia rochedoensis sp. nov. is registered only for Belmont island14 of of 40 the São Pedro e São Paulo Archipelago. The specimens were collected from seabird guano deposits over rocks. For the congeners registered in Brazil, see Figure 11.

Figure 10. Comparasion between a1, p1 and S-chaetae of abdominal segment V in: (a) Friesea rochedoensis sp. nov. (b) Friesea noronhaensis sp. nov. (c) Friesea cubensis (modified from [6]). Distribution. Friesia rochedoensis sp. nov. is registered only for Belmont island of the FigureFigure 1 10.0. ComparasionComparasion betweenSão between Pedro a1, p1 and a1,e São S-chaetae p1 Paulo and ofS Archipelago.- abdominalchaetae of segment abdominal The Vspecimens in: (a) Frieseasegment were rochedoensis Vcollected in: sp.(a) nov. Frieseafrom (b )seabirdFriesea guano rochedoensisnoronhaensis sp.sp. nov. nov (c.) (Frieseab) depositsFriesea cubensis noronhaensis over(modiﬁed rocks. from For sp. [ 6the ]).nov. congeners (c) Friesea registered cubensis in (modified Brazil, see fromFigure [6]). 11. Distribution. Friesia rochedoensis sp. nov. is registered only for Belmont island of the São Pedro e São Paulo Archipelago. The specimens were collected from seabird guano deposits over rocks. For the congeners registered in Brazil, see Figure 11.

FigureFigure 11.11. DistributionDistribution mapmap forfor Brazilian Brazilian species species of of the the genus genusFriesea FrieseaDalla Dalla Torre, Torre, 1895. 1895. Scale Scale bar: bar: 30003000 km.km.

Identification Key for the Species of Friesea Recorded in Brazil 1 Without furca...... 2 – With furca...... 7 2 1 + 1 eyes...... F. curupira Queiroz and Mendonça, 2015 – With more than 1 + 1 eyes...... 3 3 Modified chaetae of Abd VI are spiniform...... F. josei Palacios-Vargas, 1986 Figure 11. Distribution map– Modified for Brazilian chaetae species of Abd of VI the are genus clavate...... Friesea Dalla Torre, 1895. Scale bar: 4 3000 km. 4 With 10 clavate chaetae on the abd VI...... 5 – Less than 10 clavate chaetae on Abd VI...... 6 5 Th I with 3 + 3 chaetae...... F. reducta Denis, 1931 Identification Key for– the Th ISpecies with 2 + 2of ch Frieseaaetae...... Recorded F. in multiclavata Brazil Neves, Mendonça and Queiroz, 2019 1 Without furca...... 6 With 6 slightly...... clavate chaetae on abd VI...... F. boitata Queiroz and Mendonça, 2 2015 – With furca...... – With 8 clavate chaetae on Abd VI...... F. jurubatiba...... Silveira and Mendonça,...... 7 2018 7 4 + 4 eyes...... F. arlei Massoud and Bellinger, 1963 2 1 + 1 eyes...... F. curupira Queiroz and Mendonça, 2015 – With more than 1 + 1 eyes...... 3 3 Modified chaetae of Abd VI are spiniform...... F. josei Palacios-Vargas, 1986

– Modified chaetae of Abd VI are clavate...... 4 4 With 10 clavate chaetae on the abd VI...... 5 – Less than 10 clavate chaetae on Abd VI...... 6 5 Th I with 3 + 3 chaetae...... F. reducta Denis, 1931 – Th I with 2 + 2 chaetae...... F. multiclavata Neves, Mendonça and Queiroz, 2019 6 With 6 slightly clavate chaetae on abd VI...... F. boitata Queiroz and Mendonça, 2015 – With 8 clavate chaetae on Abd VI...... F. jurubatiba Silveira and Mendonça, 2018 7 4 + 4 eyes...... F. arlei Massoud and Bellinger, 1963

Insects 2021, 12, 268 15 of 39

Identification Key for the Species of Friesea Recorded in Brazil 1 Without furca...... 2 – With furca...... 7 2 1 + 1 eyes...... F. curupira Queiroz and Mendonça, 2015 – With more than 1 + 1 eyes...... 3 3 Modified chaetae of Abd VI are spiniform...... F. josei Palacios-Vargas, 1986 – Modified chaetae of Abd VI are clavate...... 4 4 With 10 clavate chaetae on the abd VI...... 5 – Less than 10 clavate chaetae on Abd VI...... 6 5 Th I with 3 + 3 chaetae...... F. reducta Denis, 1931 – Th I with 2 + 2 chaetae...... F. multiclavata Neves, Mendonça and Queiroz, 2019 6 With 6 slightly clavate chaetae on abd VI...... F. boitata Queiroz and Mendonça, 2015 – With 8 clavate chaetae on Abd VI...... F. jurubatiba Silveira and Mendonça, 2018 7 4 + 4 eyes...... F. arlei Massoud and Bellinger, 1963 – 8 + 8 eyes...... 8 8 Developed furca (state 1 according to [12])...... 9 – Reduced furca (state 2 according to [12])...... 11 9 Modified chaetae of Abd VI clavate and rugged, clavate tenent hair ...... F. noronhaensis sp. nov. – Modified chaetae of the Abd VI acuminate...... 10 10 Tenent hair acuminate...... F. cubensis Potapov and Banasko, 1985 – Tenent hair clavate...... F. rochedoensis sp. nov. 11 With clavate chaetae on Abd VI...... 12 – Without clavate chaetae on Abd VI...... F. mirabilis (Tullberg, 1871) 12 Ant IV with 5 sensilla...... 13 – Ant IV with 6 sensilla...... F. claviseta Axelson, 1900 13 Anal spine are the same size...... F. sublimis Macnamara, 1921 – Posterior anal spine smaller than anterior...... F. magnicornis Denis, 1931

Arlesia albipes (Folsom, 1927) Brazilian Oceanic Island Records. FN (3◦5208.8800 S; 32◦26013.5700 W), SF, 01 August 2012, E.C.A. Lima, A.S. Ferreira leg., CRFS# 15144–15145. Brazilian Occurrence. In Brazil, this species is widely distributed in the states PA, PE, PB, PI, MG and RJ [13] (Figure 12). In Brazilian oceanic island Arlesia albipes recorded only in Fernando de Noronha Archipelago on Fernando de Noronha Island (SF sites). The specimens were collected on superﬁcial soil and leaf litter. Biogeographic Distribution. Type locality of Arlesia albipes is Margarita Swamp, Canal Zone, collected by J. Zetek and I. Molino on June 28, 1923, in a termite mound near the base of a tree stump and deposited at U.S.N.M. # 40383. According to [1], this species has been found in the following biogeographic regions: Caribbean mainland (24a), Antillean and South Florida (24b), Venezuela and Guyana (25), Amazon (26), Northeast and Central Brazilian (27) and Andean (28). Neotropical species. Remarks. The specimens found in the Fernando de Noronha Archipelago are in accordance with the original description [14] and updates [15,16]. Lima and Zeppelini [3] recorded as A. sp. nov. For comparisons between Arlesia species recorded in Brazil, see Table4. Insects 2021, 12, x FOR PEER REVIEW 15 of 40

– 8 + 8 eyes...... 8 8 Developed furca (state 1 according to [12])...... 9 – Reduced furca (state 2 according to [12])...... 11 9 Modified ch a etae of Abd VI clavate and rugged, clavate tenent hair...... F. noronhaensis sp. nov. – Modified chaetae of the Abd VI acuminate...... 10 10 Tenent hair acuminate...... F. cubensis Potapov and Banasko, 1985 – Tenent hair clavate...... F. rochedoensis sp. nov. 11 With clavate chaetae on Abd VI...... 12 – Without clavate chaetae on Abd VI...... F. mirabilis (Tullberg, 1871) 12 Ant IV with 5 sensilla...... 13 – Ant IV with 6 sensilla...... F. claviseta Axelson, 1900 13 Anal spine are the same size...... F. sublimis Macnamara, 1921 – Posterior anal spine smaller than anterior...... F. magnicornis Denis, 1931

Arlesia albipes (Folsom, 1927) Brazilian Oceanic Island Records. FN (3°52′8.88″ S; 32°26′13.57″ W), SF, 01 August 2012, E.C.A. Lima, A.S. Ferreira leg., CRFS# 15144–15145. Brazilian Occurrence. In Brazil, this species is widely distributed in the states PA, PE, PB, PI, MG and RJ [13] (Figure 12). In Brazilian oceanic island Arlesia albipes recorded only Insects 2021, 12, 268 in Fernando de Noronha Archipelago on Fernando de Noronha Island (SF16 sites). of 39 The specimens were collected on superficial soil and leaf litter.

FigureFigure 12. 12Distribution. Distribution map map for Arlesia for Arlesia albipes albipes(Folsom, (Folsom, 1927) records 1927) in records Brazil. Blue in Brazil. circle, Fernando Blue circle, Fernando dede Noronha; Noronha; white white circle, circle, Brazilian Brazilian states. states. Scale bar: Scale 3000 bar: km. 3000 km. Table 4. Morphological characters of Arlesia Handschin, 1942 species recorded from Brazil. BD—body length in mm; AP—apical bulb vesicles; SF—sensoryBiogeographic formula; MT—mandibular Distribution teeth;. Type TL—type locality locality. of Arlesia albipes is Margarita Swamp, Canal Zone, collected by J. Zetek and I. Molino on June 28, 1923, in a termite mound near the Species Eyes BD AP Th I Tita I-III Color SF MT TL base of a tree stump and deposited at U.S.N.M. # 40383. According to [1], this species has A. albipes (Folsom, 1927) been5 + 5 found0.75–1.5 in the 3 following 2 + 2 biogeographic 19,19,18 Dark regions: blue Caribbean022/11111 mainland4 Panama (24a), Antillean A. arleana Mendonça and Black and 5 + 5 1.88 3 3 + 3 19,19,18 022/11111 6 Brazil Fernandes, 1999 and South Florida (24b), Venezuela and Guyanayellow (25), Amazon (26), Northeast and Central A. intermedia Fernandes and Brazilian5 + 5 0.98–1.6 (27) and 3 Andean 3 + 3 (28). 19,19,18 Neotropical Light species. gray 022/11111 5 Brazil Mendonça, 2004 Remarks. The specimens found in the Fernando de Noronha Archipelago are in accordance with the original description [14] and updates [15,16]. Lima and Zeppelini [3] Identiﬁcation Key for the Species of Arlesia Handschin, 1942 Recorded in Brazil 1 Mandible with 4 teeth ...... A. albipes (Folsom, 1927) – Mandible with more than 4 teeth ...... 2 2 Black with yellow strips, mandible with 6 teeth ...... A. arleana Mendonça and Fernandes, 1999 – Light gray, mandible with 5 teeth ...... A. intermedia Fernandes and Mendonça, 2004

3.1.2. Brachystomellidae Stach, 1949 Brachystomella agrosa Wray, 1953 Brazilian Oceanic Island Records. FN (3◦50054.2100 S; 32◦25045.5600 O), TF, 20 Jul 2012, E.C.A. Lima and D. Zeppelini leg., CRFS# 14400–14401, 14409–14414, 14422. FN (3◦50058.4800 S; 32◦2603.8300 O), SF, 25 July 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 14415–14418. FN (3◦51014.3200 S; 32◦26012.6200 O), TF, 25 July 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 14450–14455, 14457–14461, 15094–15096, 15098. FN (3◦50018.6600 S; 32◦2403.0900 O), SF, 07 August 2012, E.C.A. Lima and D.D. Silva CRFS# 14456. FN (3◦5103.7300 S; 32◦2600.3800 O), SF, 27 July 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 14399, 14419–14421, 14423, 14426–14427. FN (3◦50036.76” S; 32◦25012.64” O), SF, 26 Jul 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 9565–9568, 9593, 14449. FN (3◦50048.92” S; 32◦25013.6100 O), TF, 26 July Insects 2021, 12, x FOR PEER REVIEW 17 of 40

Table 5. Morphological characters of Brachystomella Agren, 1903 species recorded from Brazil.; AP––apical bulb vesicles; PAO–– PAO vesicleInsects numbe2021, 12r;, 268Mx T––maxilla teeth; Ten h––tenent hair; TT–– tenaculum teeth; DC––dental chaetae number; TL17–– oftype 39 locality; “?” ––missing date

Species Eyes AP PAO Mx T Th I Ten h TT Tita I–III DC TL 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 9562–9564, 9571–9592. FN (3◦48’45.0600 S; B. septemoculata Denis,32◦ 231931014.07 00 O), TF, 197 Jul+ 7 2012,1 E.C.A.5 Lima8 and2 A.S.+ 2 Ferreiraacumin leg.,ate CRFS# 3 15063,19,19,18 15069. FN5 Costa Rica B. garayae Queiroz and Weiner,(3◦5208.88” 2011 S; 32 ◦26013.572 + 002 O),3 SF, 015– August6 8– 2012,9 2 E.C.A.+ 2 acuminate Lima and A.S. 3 Ferreira 19,19,18 leg., CRFS# 5 Brazil ◦ 0 00 ◦ 0 00 B. platensis Najt and Massoud,14402–14403, 1974 14405, 14407,8 + 8 14425,2–3 14429,5 14437,7 14440.2 + 2 FNcapitate (3 52 8.88 S;3 32 2619,19,1813.57 O),5 SF, Argentina ◦ 0 00 ◦ 0 00 B. parvula (Schäffer,14 1896) Aug 2012, D.D. Silva8 + leg, 8 CRFS#3 6 14408.–8 FN7 (3351 + 350.41 capitateS; 32 26 5.44? O),19,19,18 TF, 01 August 5 Germany 2012, E.C.A. Lima and A.S. Ferreira leg., CRFS# 14406. FN (3◦51052.3100 S; 32◦26038.8000 O), B. agrosa Wray, 1953TF, 31 July 2012, E.C.A.8 + 8 Lima 1 and4 A.S. 7 Ferreira–10 2 + leg., 2 CRFS#acuminate 14424, 14428,3 19,19,18 14435, 14436,5 Puerto Rico B. ceciliae Fernandes and Mendonça,14438–14439, 2004 14443–14444. 8 + 8 1–2 4 8 2 + 2 capitate 3 18,18,17 6 Brazil B. nordestina Souza, Bellini and Weiner,Brazilian 2018 Occurrence 8 + 8. In3 Brazilian 4 oceanic6–8 2 island + 2 acuminatedB. agrosa recorded 3 in19,19,18 Fernando 5 de Brazil B. villalobosi Casagnau and NoronhaRapoport, Archipelago 1962 8 on + 8 Fernando 3 3 de–4 Noronha 6 and3 + 3 Rata capitate Island (SF, TF sites),3 18,18,17 the specimens 3 Brazil were collected in superﬁcial soil and leaf litter. In Brazil, this species is widely distributed B. saladaensis Weiner andand Najt, has 2001 been registered8 + in8 the3 following6 states7 2 CE, + 2 BA, acuminate PI, PE, PB, RN,3 ES,19,19,18 SP and RJ [135 ] Argentina B. contorta Denis,(Figure 1931 13). 8 + 8 1 5 7 2 + 2 acuminate 3 19,19,18 5 Costa Rica

FigureFigure 13. 13.Distribution Distribution map map for forBrachystomella Brachystomella agrosa agrosaWray, Wray, 1953 recorded 1953 recorded in Brazil. in BlueBrazil. circle, Blue circle, Fer- Fernandonando de de Noronha; Noronha; whitewhite circles, circles, Brazilian Brazilian states. states. Scale Scale bar: 3000 bar: km. 3000 km.

Biogeographic Distribution. The type locality of this species is Maricao, Puerto Rico, col- lectedIdentification by M. Capriles, Key for take the on RoadSpecies 27, atof 120Brachystomella m altitude. According Ågren, 1903 to [1], Recorded this species in has Brazil been1 2 + found 2 eyes……….…………….………………………. in the following biogeographic regions: Caribbean B. mainlandgarayae Queiroz (24a), Antillean and Weiner, 2011 and– With South more Florida than (24b), 2 Venezuela+ 2 eyes………………………….………………………………...….. and Guiana (25), Amazon (26), Northeast and Central 2 Brazil2 7 + (27).7 eyes……………………………………………………... Neotropical species. B. septemoculata Denis, 1931 Remarks. In addition to the original description [17], several authors contributed to – 8 + 8 eyes…………………………………………………………………………………… 3 the description morphological of this taxon, such as: [18–21]. The specimens of B. agrosa examined3 With 3 agreedental with cha theetae……………………… above descriptions. For comparisonsB. villalobosi between CassagnauBrachystomella and Rapoport, 1962 species– With recorded more than in Brazil, 3 dental see Table ch5.aetae……………………………………………………….. 4 4 With 5 dental chaetae……………………………………………………………………. 5 – With 6 dental chaetae………………………... B. ceciliae Fernandes and Mendonça, 2004 5 Th I with 2 + 2……………………………………………………………………………… 6 – Th I with 3 + 3………………………………………………….. B. parvula (Schäffer, 1896) 6 tenent hair acuminate……………………………………...………………..………….. 7 – tenent hair capitated……………………………….. B. platensis Najt and Massoud, 1974 7 Apical bulb simple……………………………………………………………………….. 8 – Apical bulb with 3 vesicles……………………………………………………………….. 9

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Table 5. Morphological characters of Brachystomella Agren, 1903 species recorded from Brazil.; AP—apical bulb vesicles; PAO— PAO vesicle number; Mx T—maxilla teeth; Ten h—tenent hair; TT— tenaculum teeth; DC—dental chaetae number; TL—type locality; “?” —missing date

Species Eyes AP PAO Mx T Th I Ten h TT Tita I–III DC TL B. septemoculata Denis, 1931 7 + 7 1 5 8 2 + 2 acuminate 3 19,19,18 5 Costa Rica B. garayae Queiroz and Weiner, 2011 2 + 2 3 5–6 8–9 2 + 2 acuminate 3 19,19,18 5 Brazil B. platensis Najt and Massoud, 1974 8 + 8 2–3 5 7 2 + 2 capitate 3 19,19,18 5 Argentina B. parvula (Schäffer, 1896) 8 + 8 3 6–8 7 3 + 3 capitate ? 19,19,18 5 Germany B. agrosa Wray, 1953 8 + 8 1 4 7–10 2 + 2 acuminate 3 19,19,18 5 Puerto Rico B. ceciliae Fernandes and Mendonça, 2004 8 + 8 1–2 4 8 2 + 2 capitate 3 18,18,17 6 Brazil B. nordestina Souza, Bellini and Weiner, 2018 8 + 8 3 4 6–8 2 + 2 acuminated 3 19,19,18 5 Brazil B. villalobosi Casagnau and Rapoport, 1962 8 + 8 3 3–4 6 3 + 3 capitate 3 18,18,17 3 Brazil B. saladaensis Weiner and Najt, 2001 8 + 8 3 6 7 2 + 2 acuminate 3 19,19,18 5 Argentina B. contorta Denis, 1931 8 + 8 1 5 7 2 + 2 acuminate 3 19,19,18 5 Costa Rica

Identiﬁcation Key for the Species of Brachystomella Ågren, 1903 Recorded in Brazil 1 2 + 2 eyes ...... B. garayae Queiroz and Weiner, 2011 – With more than 2 + 2 eyes ...... 2 2 7 + 7 eyes ...... B. septemoculata Denis, 1931 – 8 + 8 eyes ...... 3 3 With 3 dental chaetae ...... B. villalobosi Cassagnau and Rapoport, 1962 – With more than 3 dental chaetae ...... 4 4 With 5 dental chaetae ...... 5 – With 6 dental chaetae ...... B. ceciliae Fernandes and Mendonça, 2004 5 Th I with 2 + 2 ...... 6 – Th I with 3 + 3 ...... B. parvula (Schäffer, 1896) 6 tenent hair acuminate ...... 7 – tenent hair capitated ...... B. platensis Najt and Massoud, 1974 7 Apical bulb simple ...... 8 – Apical bulb with 3 vesicles ...... 9 8 PAO with 5 vesicles ...... B. contorta Denis, 1931 – PAO with 4 vesicles ...... B. agrosa Ågren, 1903 9 PAO with 6 vesicles ...... B. saladaensis Weiner and Najt, 2001 – PAO with 4 vesicles ...... B. nordestina Souza, Bellini and Weiner, 2018

3.1.3. Hypogastruridae Börner, 1906 Acherontiella sp. Examined material. Female on slide, Brazil, Fernando de Noronha Archipelago, Fer- nando de Noronha Island, (3◦50036.7600 S; 32◦25012.6400 O), SF, 26 July 2012, E.C.A. Lima and A.S. Ferreira leg., deposited at the CRFS-UEPB # 14568. Diagnosis. Color in ethanol white. Tegumental granulation rather strong. Body length: 746 µm; measurements as in Table4. Ant IV with 5 globular S-chaetae which 2 lateroexternal very large, placed in cavities, and the other 3 are smaller; dorsoexternal microsensillum and small subapical organite; present small non-retractile apical vesicle in ventro subapical position. Ant III and IV fused dorsally, the ventral separation well-marked. Sensory organ of Ant III consisting of two small internal sensilla, two sub-cylindrical (ring-shaped) guard sensilla and ventral microsensillum. Ant II–I with 11 and 7 chaetae, respectively. Eyes and PAO absent. Buccal cone typical of the genus. Pre-labral/labrum formula 4/554 chaetae, anterior labral chaetae spiniform. Dorsal chaetotaxy with rather short, ordinary chaetae, with long thin S-chaetae, their formula per half tergum: 022/11111. Abd VI without anal spines. Ventral tube with 4 + 4 chaetae. Tita I, II and III with 18, 18 and 17 chaetae, respectively. Femora I, II and III with 11, 11,10 chaetae, respectively, trochanters with 3,4,4 chaetae each, coxae I, II and III with 3, 8 and 7 chaetae. Remarks. Acherontiella sp. [3] is similar to Acherontiella candida (Delamare Deboutteville, 1952), A. colotlipana Palacios-Vargas and Thibaud, 1985 and A. kowalskiorum Weiner and Insects 2021, 12, 268 19 of 39

Najt, 1998 by type of guard sensilla of the Ant III organ (ring-shaped) and number of sensilla in Ant IV (with 5 globular sensilla). Acherontiella sp. differs from these species by anterior labrum chaetae (spiniform versus thin and smooth). For comparisons between Acherontiela species recorded in Brazil, see Table6. Distribution. Acherontiella sp. was found only on Fernando de Noronha Island in the SF region. The specimen was collected in soil litter. Additional material is required in order to properly describe this potentially new species. For other congeners records in Brazil, see Figure 14.

Table 6. Morphological characters of Acherontiella Absolon, 1913 species recorded from Brazil. S VI—spine on Abd VI; AL—anterior labral chaetae; Ten h—tenent hair; BD—body length in µm; NS—sensilla number of Ant IV; Tr—trochanter; TL—type locality; “?” —missing date

Species S VI AL Ten h BD NS Th I Tita I–III Femur Tr TL A. globulata Thibaud e Massoud, 1980 Absent Smooth 1 520 6 3 + 3 ? ? ? Guadalupe A. colotiplana Insects 2021, 12, x Palacios-VargasFOR PEER REVIEW and Thibaud, 1 450 5 3 + 3 17,17,16 12,11,10 3,4,4 Mexico 19 of 40 1985 Absent Smooth A. sp. Lima and Zeppelini, 2015 Absent Spine 1 746 5 3 + 3 18,18,17 11,11,10 3,4,4 Brazil

FigureFigure 14. 14.Distribution Distribution map map for forAcherontiella AcherontiellaAbsolon, Absolon, 1913 species1913 species recorded recorded in Brazil. in ScaleBrazil. Scale bar: bar:3000 3000 km. km.

Identiﬁcation Key for the Species of Acherontiella Recorded in Brazil Identification Key for the Species of Acherontiella Recorded in Brazil 1 Ant IV with 6 globular sensilla ...... A. globulata Thibaud e Massoud, 1980 -1 Ant Ant IV withIV with 5 globular 6 globular sensilla ...... sensilla…………….. A. globulata Thibaud e Massoud,2 1980 2- LabrumAnt IV withwith 4 smooth5 globular anterior sensilla……………….………...……………………………….. chaetae ...... 2 ...... 2 Labrum with 4 smooth anterior A.ch colotiplanaaetae………………………………………………….Palacios-Vargas and Thibaud, 1985 -………………………………………… Labrum with 4 spiniform anterior chaetae ...... A. colotiplanaA. sp.Palacios Lima and-Vargas Zeppelini, and 2015 Thibaud, 1985 Paraxenylla- Labrum zeliae withsp. 4 novspiniform Lima and anterior Zeppelini chaetae…….……. A. sp. Lima and Zeppelini, 2015 ParaxenyllaType material zeliae. sp. Holotype nov Lima male and on slide,Zeppelini Brazil, Atol das Rocas, Cemitério Island, ◦ 0 00 ◦ 0 00 (3 51 47.15Type materialS; 33 48 .54.14 HolotypeO), Oct male 2015, E.C.A. on slide, Lima Brazil, leg., deposited Atol das at the Rocas, CRFS-UEPB Cemitério # Island, (3°51′47.15″ S; 33°48′54.14″ O), Oct 2015, E.C.A. Lima leg., deposited at the CRFS-UEPB # 14567. Paratypes: 2 females on slides, Brazil, Atol das Rocas, Farol Island, (3°51′28.14″ S; 33°48′44.87″ O), guano, Sept 2015, E.C.A. Lima leg., deposited at the CRFS-UEPB # 14565, 14566. Description. Habitus typical of the genus. Color in ethanol blue. Body length average: 690 µm (n = 10); holotype measurements as in Table 2. Body with short acuminate chaetae (7–9 µm). Macrochaetae and S-chaetae (14–18 µm) about twice the length of ordinary chaetae. Antenna (Figure 15A). Ant IV with 4 thick S-chaetae and a small S-microchaeta between the 3 S-chaetae of the external group. Subapical organ present (Figure 15A). Apical bulb simple. Sensory organ of Ant III with 5 S-chaetae (2 small, S3 and S4, behind a fold of the integument, 2 longer guard chaetae S2 and S5, and one external microchaeta S1). Ant II–I with 11 and 7 chaetae, respectively. Head. Eyes 5 + 5 well pigmented. Labral formula: 27/554 chaetae. Mandible with 4 apical teeth (Figure 15B). Internal process of labium long, labial palps as in Figure 15B. Maxillae with 6 hypertrophied fringed lamellae

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14567. Paratypes: 2 females on slides, Brazil, Atol das Rocas, Farol Island, (3◦51028.1400 S; 33◦48044.8700 O), guano, Sept 2015, E.C.A. Lima leg., deposited at the CRFS-UEPB # 14565, 14566. Description. Habitus typical of the genus. Color in ethanol blue. Body length average: 690 µm (n = 10); holotype measurements as in Table2. Body with short acuminate chaetae (7–9 µm). Macrochaetae and S-chaetae (14–18 µm) about twice the length of ordinary chaetae. Antenna (Figure 15A). Ant IV with 4 thick S-chaetae and a small S-microchaeta between the 3 S-chaetae of the external group. Subapical organ present (Figure 15A). Apical bulb simple. Sensory organ of Ant III with 5 S-chaetae (2 small, S3 and S4, behind a fold of Insects 2021, 12, x FOR PEER REVIEW 20 of 40 the integument, 2 longer guard chaetae S2 and S5, and one external microchaeta S1). Ant II–I with 11 and 7 chaetae, respectively.

Figure 1Figure5. Paraxenylla 15. Paraxenylla zeliae zeliae sp. sp.nov nov.. (A ()A, ),dorsal dorsal view of of antenna; antenna; (B), (B labial), labial palp andpalp mandible and mandible apex, apex, nomenclaturenomenclature afterafter [22 [2].2]. Scale Scale bars: bars: 10 µ m.10 µm.

ThoraxHead. I with Eyes 3 + 5 3 + to 5 well4 + 4 pigmented. dorsal cha Labraletae. Dorsal formula: chaetotaxy 27/554 chaetae. of head Mandible and Th with in Figure 4 apical teeth (Figure 15B). Internal process of labium long, labial palps as in Figure 15B. 16a; m1Maxillae absent, with m2 6present hypertrophied on Th fringedII–III as lamellae. macrochaetae, ms on Th II not visualized. DorsalThorax chaetotaxy I with of 3 +abdomen 3 to 4 + 4 as dorsal in Figure chaetae. 16b Dorsal. Axial chaetotaxy S chaetae of 44/44,443; head and Thm2 in on Abd I–III andFigure a1 on 16 aAbd; m1 absent,V as macrochaetae. m2 present on Th Abd II–III VI as macrochaetae,dorsally with ms 5 on+ 5 Th chaetae II not visualized., one posterior chaetae andDorsal 3 An chaetotaxy microchaetae of abdomen on the as supra in Figure-anal 16b. valve. Axial S chaetae 44/44,443; m2 on Abd I–III and a1 on Abd V as macrochaetae. Abd VI dorsally with 5 + 5 chaetae, one posterior chaetae and 3 An microchaetae on the supra-anal valve. Leg. Subcoxa 1 I–III with 1,3,3 chaetae. Subcoxa 2 I–III with 0,1,1 chaetae. Coxae I–III with 3,6, 7 chaetae. Trochanters with 5 chaetae each, femora I, II, III with 12, 12, 11 chaetae, respectively. On each femur, 1 ventral chaeta longer than others. Tita I, II, III with 19, 19, 18 chaetae, respectively, each with 4 capitate tenent hairs, 2 internal longer than 2 external. Unguis without teeth, no empodial appendix (Figure 17a). Ventral tube with 1 + 1 chaetae. Abd II with 1 + 1 ventral chaetae, Abd III with 5 + 5 chaetae. Tenaculum with 3 teeth on each ramus. Mucrodens with two chaetae (Figure 17b). Female with 3 + 3 pregenital chaetae, 4–6 circumgenital and 2 eugenital. Male with 3 + 3 pregenital chaetae, 8–10 circumgenital and 4 + 4 eugenital. Each anal valve with 11–13 chaetae. Anus terminal, no anal spines.

Figure 16. Paraxenylla zeliae sp. nov. (a) dorsal view of head and Th I–III, scale bar: 10 µm. (b) dorsal view of Abd I–VI, scale bar: 10 µm.

Insects 2021, 12, x FOR PEER REVIEW 20 of 40

Figure 15. Paraxenylla zeliae sp. nov. (A), dorsal view of antenna; (B), labial palp and mandible apex, nomenclature after [22]. Scale bars: 10 µm. Insects 2021, 12, 268 21 of 39 Thorax I with 3 + 3 to 4 + 4 dorsal chaetae. Dorsal chaetotaxy of head and Th in Figure 16a; m1 absent, m2 present on Th II–III as macrochaetae, ms on Th II not visualized. Dorsal chaetotaxyEtymology. The of abdomen species is named as in Figure after Head 16b of. Axial the Biological S chaetae Reserve 44/44,443; Atol das m2 Rocas on Abd Mrs. Maurizélia de Brito Silva, in honor of a lifetime dedication to conservation and assist I–III and scientiﬁca1 on Abd research V as on macrochaetae. the Atol das Rocas. Abd VI dorsally with 5 + 5 chaetae, one posterior chaetae and 3 An microchaetae on the supra-anal valve.

Figure 16.Figure Paraxenylla 16. Paraxenylla zeliae sp. zeliae novsp.. ( nov.a) dorsal (a) dorsal view view of ofhead head and and Th I–III,I–III scale, scale bar: bar 10: µ10m. µm. (b) dorsal(b) dorsal view ofview Abd of I Abd–VI, I–VI, scale scale bar: bar: 10 10µm.µm.

Remarks. Paraxenylla zeliae sp. nov. is similar to P. afﬁniformis (Stach, 1930) (cosmopolitan), P. piloua Thibaud and Weiner, 1997 (New Caledonia), P. lapazana Palacios-Vargas and Vázquez, 1988 (Mexico), P. cubana Palacios-Vargas and Janssens, 2006 (Cuba), P. peruensis Palacios-Vargas and Janssens, 2006 (Peru) P. mahahualana Palacios-Vargas and Vázquez, 2018 (Mexico) by ventral tube with 1 + 1 chaetae, Th II–III m1 absent and m2 present, Abd I–III p2 and Abd V a1 as macrochaetae, Abd V S-chaetae in p4 position. The new species differ from P. cubana by tenaculum teeth (3 + 3 versus 2 + 2) and capitate tenent hair on Tita III (4 versus acuminate). Paraxenylla zeliae sp. nov. has 4,4,4 clavate tenent hair while P. lapazana 3,3,3 clavate or acuminate, P. afﬁniformis 5,5,5 acuminate and P. mahahualana 5,5,5 clavate. The new species share the same number of tenent hair with P. piloua and P. peruensis. However, P. zeliae sp. nov. differ from P. piloua by tenent hair morphology (clavate versus acuminate), body size (490–720 versus 360–400) and femora III (11 versus 10 chaetae); differ from P. peruensis by Th I dorsal chaetotaxy (4 + 4 versus 5 + 5), Abd II Insects 2021, 12, x FOR PEER REVIEW 21 of 40

Leg. Subcoxa 1 I–III with 1,3,3 chaetae. Subcoxa 2 I–III with 0,1,1 chaetae. Coxae I–III with 3,6, 7 chaetae. Trochanters with 5 chaetae each, femora I, II, III with 12, 12, 11 chaetae, Insects 2021, 12, 268 respectively. On each femur, 1 ventral chaeta longer than others. Tita I, II, III with 19, 19,22 of 39 18 chaetae, respectively, each with 4 capitate tenent hairs, 2 internal longer than 2 external. Unguis without teeth, no empodial appendix (Figure 17a). Ventral tube with 1 + 1 chaetae. Abd II with 1 + 1 ventral chaetae, Abd III with 5 + 5 chaetae.ventral (1 Tenaculum + 1 versus with 3 + 3) 3and teeth axial on chaetae each ramus. formula Mucrodens (44/44443 with versus two 44/44462). chaetae (Figure For other 1morphological7b). Female with comparisons 3 + 3 pregenital between chaetae,Paraxenylla 4–6 circumgenitalspecies recorded and 2 eugenital. in Brazil, Male see Table with 7. 3 + 3 pregenitalDistribution cha. Theetae, new 8–10 species circumgenital was found and on4 + Atol 4 eugenital. das Rocas Each Island anal invalve the with SF sites. 11– The 13specimen chaetae. was Anus collected terminal, in no sandy anal soilspines. near the psammophile vegetation and seabird guano on rocks. For the congeners registered in Brazil, see Figure 18.

Figure 1 17.7. ParaxenyllaParaxenylla zeliae zeliae sp.sp. nov nov.. (a) ( aTita) Tita III, III, scale scale bar: bar: 20 µm. 20 µ (m.b) mucrodens (b) mucrodens,, scale scale bar: 20 bar: µm. 20 µm. Etymology. The species is named after Head of the Biological Reserve Atol das Rocas Table 7. Morphological characters of Paraxenylla Murphy, 1965 species recorded from Brazil. VT—ventral tube chaetae; Mrs. Maurizélia de Brito Silva, in honor of a lifetime dedication to conservation and assist µ TT—tenaculum teeth; VC—ventralscientific chaetae research Abd on II; the Ten Atol h—tenent das Rocas. hair; BD—body length in m; NS—sensilla number of At. IV; TL—type locality. Ac—acuminate;Remarks Cl—clavate.. Paraxenylla zeliae sp. nov. is similar to P. affiniformis (Stach, 1930) (cosmopol- Species VT TTitan), P. VC piloua ThibaudTen h andBD Weiner, 1997Th I(New TitaCaledonia), I-III FemurP. lapazanaTrochanter Palacios-VargasTL and Vázquez, 1988 (Mexico), P. cubana Palacios-Vargas and Janssens, 2006 (Cuba), P. pe- Paraxenylla piloua ruensis Palacios-Vargas and Janssens, 2006 (Peru) P. mahahualana Palacios-Vargas andNew Thibaud and 1 + 1 3 1 + 1 4 Ac 360–400 3 + 3 19,19,18 12,12,10 5,5,5 Caledonia Weiner, 1997 Vázquez, 2018 (Mexico) by ventral tube with 1 + 1 chaetae, Th II–III m1 absent and m2 Paraxenylla sooretamensis present, Abd I–III p2 and Abd V a1 as macrochaetae, Abd V S-chaetae in p4 position. The 4 + 4 3 ? 0 440–820 3 + 3 19,19,18 12,12,10 5,5,4 Brazil Queiroz and new species differ from P. cubana by tenaculum teeth (3 + 3 versus 2 + 2) and capitate Deharveng, 2008 Paraxenylla zeliae tenent hair on Tita III (4 versus acuminate). Paraxenylla zeliae sp. nov. has 4,4,4 clavate 1 + 1 3 1 + 1 490–720 3 + 3 or 19,19,18 12,12,11 5,5,5 sp. nov. tenent hair while4 P. Cl lapazana 3,3,3 clavate4 + or 4 acuminate, P. affiniformis 5,5,5 acuminate Braziland P. mahahualana 5,5,5 clavate. The new species share the same number of tenent hair with P. piloua and P. peruensis. However, P. zeliae sp. nov. differ from P. piloua by tenent hair Identiﬁcation Key for Paraxenylla Murphy, 1965 Species Recorded in Brazil morphology (clavate versus acuminate), body size (490–720 versus 360–400) and femora III1 Ventral (11 versus tube 10 1 ch +a 1etae); chaetae...... differ from P. peruensis by Th I dorsal chaetotaxy (4 + 4 versus 5 2 –+ Ventral 5), Abd tube II ventral 4 + 4 chaetae...... (1 + 1 versus 3 + 3) andP. sooretamensis axial chaetaeQueiroz formula and (44/44443 Deharveng, versus 2008 44/44462).2 tenent hair For clavate,other morphological femora III with comparisons 11 chaetae...... between Paraxenylla speciesP. recorded zeliae sp. in nov. –Brazil tenent, see hair Table acuminate, 7. femora III with 10 chaetae. P. piloua Thibaud and Weiner, 1997

Willemia insularum sp. nov. Lima and Zeppelini Type material. Holotype female on the slide, Brazil, Rio Grande do Norte, Atol das Rocas, (03◦5105000 S, 33◦4804800 W), intertidal sand, 08–18 September 2015, E.C.A. Lima leg., deposited at the CRFS-UEPB # 12261. Paratypes: 2 males and 3 females on slides, same data as holotype, deposited at the CRFS-UEPB # 12262, 12263, 12264, 12265, 12266. Additional material: 2 females and 1 male on slides, Brazil, Pernambuco, Fernando de Noronha (3◦50058.4800 S; 32◦2603.8300 O and 3◦50036.7600 S; 32◦25012.6400 O), 25–26 July 2012, E.C.A. Lima and A.S. Ferreira leg., deposited at the CRFS-UEPB # 14562, 14563, 14564. Insects 2021, 12, x FOR PEER REVIEW 22 of 40

Table 7. Morphological characters of Paraxenylla Murphy, 1965 species recorded from Brazil. VT––ventral tube chaetae; TT–– tenaculum teeth; VC––ventral chaetae Abd II; Ten h––tenent hair; BD––body length in µm; NS––sensilla number of At. IV; TL– –type locality. Ac––acuminate; Cl––clavate.

Species VT TT VC Ten h BD Th I Tita I-III Femur Trochanter TL Paraxenylla piloua Thibaud New Cale- 1 + 1 3 1 + 1 4 Ac 360–400 3 + 3 19,19,18 12,12,10 5,5,5 and Weiner, 1997 donia Paraxenylla sooretamensis Quei- 4 + 4 3 ? 0 440–820 3 + 3 19,19,18 12,12,10 5,5,4 Brazil roz and Deharveng, 2008 3 + 3 or Paraxenylla zeliae sp. nov. 1 + 1 3 1 + 1 4 Cl 490–720 19,19,18 12,12,11 5,5,5 Brazil 4 + 4

Distribution. The new species was found on Atol das Rocas Island in the SF sites. The Insects 2021, 12, 268 23 of 39 specimen was collected in sandy soil near the psammophile vegetation and seabird guano on rocks. For the congeners registered in Brazil, see Figure 18.

FigureFigure 18. 18Distribution. Distribution map map for Paraxenylla for ParaxenyllaMurphy, Murphy, 1965 species 1965 recorded species in Brazil.recorded Scale in bar: Brazil. 3000 kmScale. bar: 3000 km. Description. Habitus typical of the genus. Color in ethanol white. Tegumental gran- µ n ulationIdentification ﬁne and Key regular. for Paraxenylla Body length: Murphy, 490 m 1965 ( = Species 5); holotype Recorded measurements in Brazilas in Table2. 1 VentralBody with tube short 1 + acuminate1 chaetae...... ordinary chaetae, some slightly longer. Sensory chaetae .... 2 are– Ventral lanceolate tube and 4 + longer 4 cha thanetae...... ordinary chaetae... P. Antennaesooretamensis are somewhat Queiroz shorterand Deharveng, than 2008 cephalic2 tenent diagonal. hair clavate, femora III with 11 chaetae...... P. zeliae sp. nov. – tenentAntenna hair (Figure acuminate, 19). Ant femora IV with III 4 (S1,with S2, 10 S8, ch S9)aetae. subcylindrical P. piloua sensilla,Thibaud 2 globularand Weiner, 1997 sensilla S7 and S4, hidden each in a separate cavity. Microsensillum (ms) set in the cavity ofWillemia S7 sensillum. insularum Sensilla sp. S2nov. and Lima S9 hardly and Zeppelini differentiated from ordinary chaetae, S1 and S8 subcylindrical.Type material Apical. Holotype bulb small female and on oval, the subapical slide, Brazil, organelle Rio setGrande in a deep do Norte, cavity, Atol das roundish. Sensorial organ of Ant III with 2 long guard chaetae, 2 long subcylindrical Rocas, (03°51′50″ S, 33°48′48″ W), intertidal sand, 08–18 September 2015, E.C.A. Lima leg., sensilla, 1 ventral microsensillum, and a large integumentary fold hiding 2 internal sensilla. Antdeposited II and I withat the 11 CRFS and 6- chaetae,UEPB # respectively. 12261. Paratypes: 2 males and 3 females on slides, same dataEyes as holotype, and a0 chaetae deposited absent, PAO at the with CRFS 7–8 vesicles-UEPB ( Figure# 12262, 20a 12263,). Head 12264, dorsal (Figure 12265, 20 12266.b). Addi- Prelabrum/labrumtional material: 2 females with 2/534 and chaetae 1 male (Figure on slides, 21a). Brazil, Along cephalicPernambuco, groove Fernando 3 + 3 chaetae de Noronha (Figure(3°50′58.48 21b).″ S; 32°26′3.83″ O and 3°50′36.76″ S; 32°25′12.64″ O), 25–26 July 2012, E.C.A. LimaDorsal and A.S. chaetotaxy Ferreira (Figures leg., 20depositedb and 22). at S-chaetae the CRFS per-UEPB half tergum # 14562, formula: 14563, 22/11111, 14564. in m7 and p4 position on Th II and III, in p4 position on abdominal I to IV and p2 position on Abd V. Sensilla p4 on Th II–Abd I easily differentiated from ordinary chaetae. Abd IV without m-row. With two anal spines on m1 position. Ventral abdominal chaetotaxy is presented in Figure 23a. Sternum of Abd IV with 12 + 12 chaetae. Sternum of Abd II with a-row with 2 chaetae on each side (a3 present). Anal lobes with 16 + 16 chaetae (2 i, 3 hr, e and b absent). Ventral tube with 4 + 4 chaetae (Figure 23b). Tita I, II and III with 12, 12, 11 chaetae. Unguis without teeth, empodial appendage vestigial (Figure 23c). Etymology. An allusion for being the only new species of this study that occurs in two different island regions. Remarks. Willemia insularum sp. nov. shares characteristics with the buddenbrocki-group species sensu D’Hese 2000 [23–25] by S-chaetae S4 and S7 of Ant IV both large, globular, placed in the cavity, and covered in part by tegumental fold and absence of a0 chaetae on Insects 2021, 12, x FOR PEER REVIEW 23 of 40

Insects 2021, 12, 268 24 of 39

Description. Habitus typical of the genus. Color in ethanol white. Tegumental granu- thelation head. fine The and new regular. species Body presents length: apical 490 vesicle, µm (n Abd= 5); Vholotype with S-chaetae measurements in p2 position; as in Table 2. 3 + 3 chaetaeBody withon Th short I, presence acuminate of anal spines;ordinary Abd ch IVaetae, and V some with S-chaetaeslightly longer. lanceolate. Sensory The chaetae neware lanceolate species is similar and longer to W. dheseithan ordinaryBu, Potapov cha andetae. Gao, Antenna 2012 (Shandong,e are somewhat Paciﬁc shorter coast than ce- ofphalic China), diagonal.W. delamarei Prabhoo, 1971 (Indian region); W. antennomonstrum Bu, Potapov and Gao,Antenna 2012 (Hainan, (Figure South 19). Ant China). IV withWillemia 4 (S1, insularum S2, S8, sp.S9) nov.subcylindrical differs from sensilla,W. dhesei 2 globular bysensilla chaetotaxy S7 and of AbdS4, hidden IV (without each m-rowin a separate versus m1cavity. to m4 Microsensillum present) and postantennal (ms) set in the cavity organ (7–8 versus 5–6 vesicles). W. delamarei has a postantennal organ with 8–12 vesicles, of S7 sensillum. Sensilla S2 and S9 hardly differentiated from ordinary chaetae, S1 and S8 S-chaetae absent in Th II and Abd I while the new species present postantennal organ with 7–8subcylindrical. vesicles and 2 A S-chaetaepical bulb in Thsmall II (m7 and and oval, p4) subapical and 1 in Abd organelle I (p4). Willemia set in a insularum deep cavity, roun- sp.dish. nov., Sensorial can be easily organ distinguished of Ant III with from 2W. long antennomonstrum guard chaetae,by 2 the long sensory subcylindrical organ of sensilla, Ant1 ventral III usual microsensillum, for the genus versus and a 2 large expanded integumentary and granulated fold guard hiding sensilla. 2 internal For other sensilla. Ant II morphologicaland I with 11 comparisons and 6 chaetae, between respectively.Willemia species recorded in Brazil, see Table8.

Figure 19. Willemia insularum sp. nov. 33, dorsal view of antenna. Scale bar: 10 µm. Figure 19. Willemia insularum sp. nov. 33, dorsal view of antenna. Scale bar: 10 µm.

Eyes and a0 chaetae absent, PAO with 7–8 vesicles (Figure 20a). Head dorsal (Figure 20b). Prelabrum/labrum with 2/534 chaetae (Figure 21a). Along cephalic groove 3 + 3 chaetae (Figure 21b).

Insects 2021, 12, 268 25 of 39

Insects 2021, 12, x FOR PEER REVIEW 24 of 40

Figure 20. Willemia insularum sp. nov. (a) postantennal organ, scale bar: 20 µm. (b) dorsal view of head and thorax I–III. Scale bar:Figure 10 µm. 20. Willemia insularum sp. nov. (a) postantennal organ, scale bar: 20 µm. (b) dorsal view of Insects 2021, 12, x FOR PEER REVIEW 25 of 40 head and thorax I–III. Scale bar: 10 µm.

Figure 21. Willemia insularum sp. nov. (a) prelabrum/labrum, scale bar: 10 µm. (b) ventral view of headFigure, scale bar: 21. 10 Willemiaµm. insularum sp. nov. (a) prelabrum/labrum, scale bar: 10 µm. (b) ventral view of head,Dorsal scale chaetotaxy bar: (Fig 10ureµsm. 20b and 22). S-chaetae per half tergum formula: 22/11111, in m7 and p4 position on Th II and III, in p4 position on abdominal I to IV and p2 position on Abd V. Sensilla p4 on Th II–Abd I easily differentiated from ordinary chaetae. Abd IV without m-row. With two anal spines on m1 position. Ventral abdominal chaetotaxy is presented in Figure 23a. Sternum of Abd IV with 12 + 12 chaetae. Sternum of Abd II with a-row with 2 chaetae on each side (a3 present). Anal lobes with 16 + 16 chaetae (2 i, 3 hr, e and b absent). Ventral tube with 4 + 4 chaetae (Figure 23b). Tita I, II and III with 12, 12, 11 chaetae. Unguis without teeth, empodial appendage vestigial (Figure 23c).

Insects 2021, 12, 268 26 of 39 Insects 2021, 12, x FOR PEER REVIEW 26 of 40

FigureFigure 22. 22.Willemia Willemia insularum insularumsp. sp. nov., nov., dorsal dorsal view view of of Abd Abd I–VI, I–VI, anal anal spine spine details. details. Scale Scale bars: bars: 10 µm 10 µm and anal spine detail, 20 µm. and anal spine detail, 20 µm.

InsectsInsects2021 2021, 12, ,12 268, x FOR PEER REVIEW 27 of27 40 of 39

Figure 23. Willemia insularum sp. nov. (a) ventral view of Abd II–VI, scale bar: 10 µm. (b) ventral µ Figuretubes, scale 23. Willemia bar: 20 µm. insularum (c) Titasp. III, nov.scale (bar:a) ventral 20 µm. view of Abd II–VI, scale bar: 10 m. (b) ventral tubes, scale bar: 20 µm. (c) Tita III, scale bar: 20 µm. Etymology. An allusion for being the only new species of this study that occurs in two Table 8. Data matrix of 52 charactersdifferent coded island for regions. Willemia species recorded in Brazil. Character numbers correspond to the characters listed in [26]. PolymorphicRemarks characters. Willemia coded insularum as follow [sp.23], nov. 0 or 1 shares = a; 0 or characteristics 2 = b; 0, 2 or 3 = with c. “?”—missing the buddenbrocki date. - group species sensu D’Hese 2000 [23–25] by S-chaetae S4 and S7 of Ant IV both large, Speciesglobular, placed in the 00000 cavity, 00,000 and 11111 covered 11,111 in 22222 part by 22,222 tegumental 33333 33,333 fold 44444 and absence 44,444 55 of a0 chaetae on the head. 01234The new 56,789 species 01234 presents 56,789 01234 apical 56,789 vesicle, 01234 Abd 56,789 V with 01234 S-ch 56,789aetae 01 in p2 position; 3 + 3 chaetae on Th I, presence of anal spines; Abd IV and V with S-chaetae lan- W. brevispina Hüther, 1962 13032 03,111 11311 11,101 00122 10,100 01101 11,111 11111 11,111 01 ceolate. The new species is similar to W. dhesei Bu, Potapov and Gao, 2012 (Shandong, W. zeppelini D’Haese and Thibaud, 2011 0c0b2 2b111 01,211 1110? 11,122 10000 01,101 11110 11,001 01111 11 W. insularum sp. nov.Pacific coast of China), 13032 W. 03,111delamarei 11?11 Prabhoo, 10,101 00033 1971 11,110(Indian 11111 region); 11,111 W. 11011 antennomonst 11,100 01 rum

Insects 2021, 12, x FOR PEER REVIEW 28 of 40

Bu, Potapov and Gao, 2012 (Hainan, South China). Willemia insularum sp. nov. differs from W. dhesei by chaetotaxy of Abd IV (without m-row versus m1 to m4 present) and postantennal organ (7–8 versus 5–6 vesicles). W. delamarei has a postantennal organ with 8–12 vesicles, S-chaetae absent in Th II and Abd I while the new species present postantennal organ with 7–8 vesicles and 2 S-chaetae in Th II (m7 and p4) and 1 in Abd I (p4). Willemia insularum sp. nov., can be easily distinguished from W. antennomonstrum by the sensory organ of Ant III usual for the genus versus 2 expanded and granulated guard sensilla. For other morphological comparisons between Willemia species recorded in Brazil, see Table 8.

Table 8. Data matrix of 52 characters coded for Willemia species recorded in Brazil. Character numbers correspond to the characters listed in [26]. Polymorphic characters coded as follow [23], 0 or 1 = a; 0 or 2 = b; 0, 2 or 3 = c. “?”—missing date.

Species 00000 00,000 11111 11,111 22222 22,222 33333 33,333 44444 44,444 55 01234 56,789 01234 56,789 01234 56,789 01234 56,789 01234 56,789 01 W. brevispina Hüther, 1962 13032 03,111 11311 11,101 00122 10,100 01101 11,111 11111 11,111 01 W. zeppelini D’Haese and Thibaud, 2011 0c0b2 2b111 01,211 1110? 11,122 10000 01,101 11110 11,001 01111 11 Insects 2021, 12, 268 28 of 39 W. insularum sp. nov. 13032 03,111 11?11 10,101 00033 11,110 11111 11,111 11011 11,100 01

DistributionDistribution. The. The new new species species were foundwere onfound Fernando on Fernando de Noronha de and Noronha Atol das and Atol das RocasRocas Islands Islands in thein the SF sites. SF sites. The specimens The specimens were collected were col in sandlected near in the sand psammophile near the psammophile vegetation,vegetation, litter litter and and seabird seabird guano. guano. For the For congeners the congeners registered inregistered Brazil, see in Figure Brazil, 24. see Figure 24.

FigureFigure 24. 24Distribution. Distribution map map for Willemia for WillemiaBörner, 1901Börner, species 1901 recorded species in recorded Brazil. Scale in bar: Brazil. 3000 km.Scale bar: 3000 km.

Identiﬁcation Key for Willemia Species Recorded in Brazil Identification Key for Willemia Species Recorded in Brazil 1 Anal spine absent ...... W. zeppelinii D’Hese and Thibaud, 2011 1 Anal spine absent……………………………... W. zeppelinii D’Hese and Thibaud, 2011 – Anal spine present ...... 2 2– AbdAnal IV andspine V with present……………………………………………………………………….. S-chaetae subcylindrical and acuminate ...... 2 ...... 2 Abd...... IV and V with S-chaetae subcylindrical andW. acuminate……...…………………. brevispina Hüther, 1962 –…………………………………………………………… Abd IV and V with S-chaetae lanceolate ...... …….W.W. insularum brevispinasp. nov. Hüther, 1962 Xenylla– Abd yucatanaIV and Mills,V with 1938 S-chaetae lanceolate…………………….... W. insularum sp. nov.

Brazilian Oceanic Island Records. FN (3◦50043.0000 S; 32◦25034.0400 W), SF, 20 Jul 2012, E.C.A Lima and D. Zeppelini leg., CRFS# 14518, 14519, 14522, 14528. FN (3◦50054.2100 S; 32◦25045.5600 W), TF, 20 July 2012, E.C.A Lima and D. Zeppelini leg., CRFS# 14520, 14525, 14527, 14529, 14530, 14531, 14535, 14536, 14553, 14560, 14561. FN (3◦50031.8500 S; 32◦24014.7400 W), TF, 07 August 2012, E.C.A Lima and D.D. Silva leg, CRFS#14474, 14491, 14545, 14546, 14547, 14548, 14549, 14550. FN (3◦50018.6600 S; 32◦2403.0900 W), SF, 07 August 2012, E.C.A Lima and D.D. Silva leg, CRFS# 14479, 14487, 14489, 14492, 14503–14505, 14511–14517, 14521, 14523–14524, 14526, 14532–14537, 14538–14544, 14551–14552, 14555–14556, 14558–14559, 14734, 14735–14768. FN (3◦5103.7300 S; 32◦2600.3800 W), SF, 27 July 2012, E.C.A Lima and A.S. Ferreira leg., CRFS# 14478, 14481, 14484. FN (3◦50036.7600 S; 32◦25012.6400 W), SF, 26 July 2012, E.C.A Lima and A.S. Ferreira leg., CRFS# 9373, 9374–9384, 9423–9533. FN (3◦50048.9200 S; 32◦25013.6100 W), TF, 26 Jul 2012, E.C.A Lima and A.S. Ferreira leg., CRFS# 9532. FN (3◦51050.4100 S; 32◦2605.4400 W), TF, 01 August 2012, E.C.A Lima and A.S. Ferreira leg., CRFS# 14462–14472, 14476–14477, 14480, 14485, 14486, 14493–14502. Brazilian Occurrence. In Brazil, this species had already been registered in the states of ES, RJ and the island of Fernando Noronha [13] (Figure 25). Cosmopolitan species. Insects 2021, 12, x FOR PEER REVIEW 30 of 40

Identification Key for Xenylla Tullberg, 1869 Species Recorded in Brazil 1 With 5 + 5 eyes……………………………………………………………………………... 2 – With 4 + 4 eyes…………………………………………………….. X. yucatana Mills, 1938 2 Ant IV with 4–5 sensilla…………………………………………………………………. 3 – Ant IV with 6 sensilla……………………………………………………....……………. 8 3 With 1,1,1 tenent hair……………………………………. X. brasiliensis da Gama, 1978 – With 1,2,2 or 2,2,2 tenent hair…………………………………………………...………. 4 4 With 1,2,2 tenent hair……………………………………………. X. welchi Folsom, 1916 – With 2,2,2 tenent hair……………………………………...…………………………….. 5 5 Manubrium with 28 chaetae…………………….... X. hodori Neves and Mendonça, 2017 – Manubrium with more than 28 chaetae…………………………………...………...... 6 6 Mucro separate from dens………………………………………………………………. 7 – Mucro and dens fused …………………………………………………………………... 9 7 Creased cuticular structure and p6 chaetae present on Abd IV……………………….. …………………………………………………...X. manuelae Queiroz and Mendonça, 2016 – Creased cuticular structure and p6 chaetae absent on Abd IV………………………... ………………..……………………………….. X. capixaba Fernandes and Mendonça, 2010 8 With 1,1,1 tenent hair………………………...………. X. nirae Gama and Oliveira, 1994 – With 1,2,2 tenent hair……………………….... X. wandae Queiroz and Mendonça, 2016 Insects 2021, 12, 268 9 Dens with 2 posterior chaetae……………………………… X. maritima29 of 39 Tullberg, 1869 – Dens with 1 posterior chaetae………...………………... X. subcavernarum Gama, 1969

FigureFigure 25. 25Distribution. Distribution map map for Xenylla for Xenylla yucatana yucatanaMills, 1938 Mills, records 1938 in Brazil. records Scale in bar: Brazil. 3000 Scale km. bar: 3000 km. BlueBlue circle, circle, Fernando Fernando de Noronha; de Noronha; white circles, white Brazilian circles, states.Brazilian Scale states. bar: 3000 Scale km. bar: 3000 km. Biogeographic Distribution. The type locality of this species in Mexico, Yucatan Penin- sula,3.1.4. San Isotogastruridae Bulha Cenote Motul; Thibaud specimens and collected Najt, 1992 in bat feces [27]. Xenylla yucatana has circumtropical distribution distributed in the following biogeographic regions African IndianIsotogastrura Desert (9), mucrospatulata East African Steppe Palacios (13), South-Vargas, African Lima (14), and Madagascar Zeppelini, (15), 2013 Ascension and St Helena (16), Continental South and East Asia (18), Malaysian (19), Hawaiian (20), New CaledoniaBrazilian (21),Oceanic Melanesia Island and Records Micronesia. FN (22), (3°50 Caribbean′40.61″ MainlandS, 32°25’ (24a),35.41 Antillean″ O), SB, 20 July 2012, andE.C.A. South Lima Florida and (24b), A.S. Venezuela Ferreira and Leg., Guyana CRFS# (25), Northeast3680–3696 and. FN Central (3°50 Brazilian′13.36″ (27),S, 32°24’0.28″ O), AndeanSB, 07 Aug (28), Northust 2012, and EastE.C.A Australia Lima (32)and according D. Zeppelini to [1]. leg., CRFS# 3697–3701. FN (3°50′30.12″ S, 32°25Remarks’3.84. The” O), specimens SB, 20 Jul analyzedy 2012 agree, E.C.A. with Lima descriptions and A.S. by [Ferreira28–31]. ForLeg., other CRFS# 3702. FN morphological(3°50′30.12″ S, comparisons 32°25′3.84 between″ O), SB,Xenylla 20 Julspecies 2012, recorded E.C.A. in Lima Brazil, and see Table A.S.9 .Ferreira In the Leg., CRFS# oceanic islands, the species was found in great abundance on the island of Fernando de Noronha in the SF and TF sites.

Table 9. Morphological characters of Xenylla Tullberg, 1869 species recorded from Brazil.; Ten h—tenent hair; OCM— ordinary chaetae morphology; TL—type locality; MD—mucrodens, MSD—mucro separate from dens; “?”—missing date.

Species Eyes Ant IV Sensilla Furca Manubrial Chaeta OCM Ten h TL X. maritima Tullberg, 1869 5 + 5 4 MD ? Smooth 2,2,2 ? X. nirae Gama and Oliveira, 1994 5 + 5 6 MSD ? ? 1,1,1 Brazil X. brasiliensis da Gama, 1978 5 + 5 4 MD ? Smooth 1,1,1 Brazil X. subcavernarum Gama, 1969 5 + 5 4 MD ? ? 2,2,2 Brazil X. welchi Folsom, 1916 5 + 5 4 MSD 34 Smooth 1,2,2 North America X. hodori Neves and Mendonça, 2017 5 + 5 4 MSD 28 Serrated 2,2,2 Brazil X. yucatana Mills, 1938 4 + 4 4 MSD 30 Smooth 2,2,2 Mexican X. manuelae Queiroz and Mendonça, 5 + 5 4 MSD 34 2,2,2 2016 Serrated Brazil X. capixaba Fernandes and 5 + 5 4–5 MSD 34 2,2,2 Mendonça, 2010 Serrated Brazil X. wandae Queiroz and Mendonça, 5 + 5 6 MSD 34 1,2,2 2016 Serrated Brazil Insects 2021, 12, 268 30 of 39

Identiﬁcation Key for Xenylla Tullberg, 1869 Species Recorded in Brazil 1 With 5 + 5 eyes ...... 2 – With 4 + 4 eyes ...... X. yucatana Mills, 1938 2 Ant IV with 4–5 sensilla ...... 3 – Ant IV with 6 sensilla ...... 8 3 With 1,1,1 tenent hair ...... X. brasiliensis da Gama, 1978 – With 1,2,2 or 2,2,2 tenent hair ...... 4 4 With 1,2,2 tenent hair ...... X. welchi Folsom, 1916 – With 2,2,2 tenent hair ...... 5 5 Manubrium with 28 chaetae ...... X. hodori Neves and Mendonça, 2017 – Manubrium with more than 28 chaetae ...... 6 6 Mucro separate from dens ...... 7 – Mucro and dens fused ...... 9 7 Creased cuticular structure and p6 chaetae present on Abd IV ...... X. manuelae Queiroz and Mendonça, 2016 – Creased cuticular structure and p6 chaetae absent on Abd IV ...... X. capixaba Fernandes and Mendonça, 2010 8 With 1,1,1 tenent hair ...... X. nirae Gama and Oliveira, 1994 – With 1,2,2 tenent hair ...... X. wandae Queiroz and Mendonça, 2016 9 Dens with 2 posterior chaetae ...... X. maritima Tullberg, 1869 – Dens with 1 posterior chaetae ...... X. subcavernarum Gama, 1969

3.1.4. Isotogastruridae Thibaud and Najt, 1992 Isotogastrura mucrospatulata Palacios-Vargas, Lima and Zeppelini, 2013 Brazilian Oceanic Island Records. FN (3◦50040.6100 S, 32◦25’35.4100 O), SB, 20 July 2012, E.C.A. Lima and A.S. Ferreira Leg., CRFS# 3680–3696. FN (3◦50013.3600 S, 32◦24’0.2800 O), SB, 07 August 2012, E.C.A Lima and D. Zeppelini leg., CRFS# 3697–3701. FN (3◦50030.1200 S, 32◦25’3.84” O), SB, 20 July 2012, E.C.A. Lima and A.S. Ferreira Leg., CRFS# 3702. FN (3◦50030.1200 S, 32◦2503.8400 O), SB, 20 Jul 2012, E.C.A. Lima and A.S. Ferreira Leg., CRFS# 3703–3708. FN (3◦51019.9800 S, 32◦26043.2300 O), SB, 20 July 2012, E.C.A. Lima and A.S. Ferreira Leg., CRFS# 3709–3710. Distribution: This species is known only in the sand beaches of the Fernando de Noronha Islands. Small body size (350 µm) of I. mucrospatulata indicates that it inhabits narrow passages among the grains of sand [32]. Interstitial Collembola looks like typical euedaphic species but is ﬂexible and slender enough to be able to move between sand grains of small size without changing the pore architecture [33]. All specimens in this study were found in the sand at the intertidal zone (SB sites). Remark. The specimens analyzed agree with the original description by [34]. For others, morphological comparisons between Isotogastrura species recorded in Brazil, see Table 10.

Table 10. Morphological characters of Isotogastrura species record from Brazil modified from [34]; 1, Abd V tegumentary tubercle; 2. Abd I sternite chaetae number; 3. Abd III tergite with a pair of tubercles; 4. Prelabral chaetae (presence vs. absence); 5. Sensillum of Ant III simple or bifid; 6. Dental chaetae; 7. Shape of mucro; 8. Genital chaetae; 9. Number of sensilla on Ant IV; 10. Tubercles on Abd VI; 11. Chaetae on labial triangle; 12. Chaetae on manubrium; 13. Chaetae on furcal subcoxa; 14. Basal manubrium Chaetae. Abbreviations: BI = bifid; H = hooked; MOD = modified; SP = spatulate; un = unmodified; “?”—missing date.

µm 1 2 3 4 5 6 7 8 9 10 11 12 13 14 I. mucrospatulata 350 + 1 — — BI 4 SP MOD 7 + 1 — 4 16 5/2 5 + 5 I. praiana 450 + ? + + BI 4 H un 6 — 5 13 5/2 ? Insects 2021, 12, 268 31 of 39

3.2. Species List of Brazilian Poduromorpha In Brazil, Poduromorpha fauna is registered in 18 states and 3 oceanic islands (Table 11). The states of Acre, Alagoas, Goiás, Maranhão, Roraima, Rio Grande do Sul, Santa Catarina, Tocantins and the oceanic Archipelago of Trindade and Martim Vaz have no record of Poduromorpha. Including the new taxa presented in this study, 139 Brazilian species of Poduromopha distributed in the families Brachystomellidae Stach, 1949 (24 spp.); Hy- pogastruridae Börner, 1906 (31 spp.); Isotogastruridae Thibaud and Najt, 1992 (2 spp.); Neanuridae Börner, 1901 (71 spp.); Odontellidae Massoud, 1967 (1 sp.); Onychiuridae Lubbock, 1867 (4 spp.); Tullbergiidae Bagnall, 1935 (6 spp.), see Table2. This number represents an increase of 41 new species records since the last synthesis by [2].

Table 11. Species of Poduromorpha recorded from Brazilian states and oceanic islands–updated from [2]. *—New record after [2]. In parentheses, information obtained from non-original reference. “?” following state abbreviation—questionable state record. “un”—unspeciﬁed or unknown Brazilian collection habitats. World distribution—see Methods/Species list of Brazilian Poduromorpha. “?” following the distribution abbreviation—questionable distribution record. Habitat— representative type of habitat for the species. “un”—unspeciﬁed or unknown habitat (Brazilian distribution, locality abbreviations refer to Brazilian states and Insular locality as listed in Table1). Family names in bold.

Family/Species Brazil Distribution World Distribution Habitats Record Brachystomellidae Stach, 1949 Halophyte–psammophyte vegetation, sand dunes, flooded areas, pasture, eucalyptus, citric Brachystomella agrosa Wray, 1953 FN, PE, BA, PE, RJ, SP Neo [3,20,35]* plantation, understory, riparian vegetation, moss, litter and soil over beach rocks, oceanic island Brachystomella aspera (Börner, 1906) São Francisco NCB? un [36] Halophyte–psammophyte Brachystomella ceciliae Fernandes NCB and Mendonça, 2004 RJ (ES) vegetation, litter on sand dunes, [37] flooded areas Halophyte–psammophyte Brachystomella contorta Denis, 1931 RJ, (ES) Bor, Neo, Pal vegetation, sand dunes, beach [37,38] sand, moss, soil over beach rocks Brachystomella garayae Queiroz and ES NCB Weiner, 2011 Forest litter [39] Brachystomella nordestina Souza, Leaf litter over sandy soil of an RN NCB [40]* Bellini and Weiner, 2018 urban forested Brachystomella parvula (Schäffer, RN Cos 1896) Forest litter [41] Beach sand, litter over sandy Brachystomella platensis Najt and ES, RJ Neo, Aus dunes, halophyte–psammophyte [42,43]* Massoud, 1974 vegetation, guriri vegetation, dunes and supralittoral vegetation Brachystomella saladaensis Weiner CE Neo and Najt, 2001 Leaf litter on the shore of lakes [44] Brachystomella septemoculata Denis, RJ 1931 Pal, Neo Dunes, rotten wood [45] Brachystomella villalobosi Casagnau PE Neo un [35] and Rapoport, 1962 Brachystomellides compositus Arlé, RJ Neo 1959 Beach sand, litter [46] Folsomiella albida (Arlé, 1959) RJ Neo Leaf litter [38,46] Folsomiella ceca (Folsom, 1927) (RJ) Neo Soil [47] Folsomiella intermedia (Arlé, 1939) RJ NCB Soil [38,48] Folsomiella pseudoceca Mendonça, RJ NCB Fernandes and Abrantes, 2005 Soil [49] Folsomiella trisetosa Mendonça, RJ, SP NCB Soil and rotten logs, soil near Fernandes and Abrantes, 2005 streams [49] Maricaella duna Mendonça and RJ NCB Fernandes, 1997 Leaf litter, sand dunes [50] Micronella itacaman Queiroz and RJ, MG NCB Mendonça, 2013 Leaf litter and soil, altitude 2400 m [51]* Micronella longisensilla Queiroz and RJ NCB Leaf litter and soil, altitude 2100 m [51]* Mendonça, 2013 Micronella porcus (Denis, 1933) RJ Neo Soil [46] Neorganella rotundatae Queiroz and RJ, MG NCB Mendonça, 2013 Leaf litter and soil, altitude 2400 m [51]* Halophyte–psammophyte Rapoportella pitomboi Mendonça MG, RJ NCB and Fernandes, 1995 vegetation, sand dunes, flooded [37,52] areas Setanodosa occidentalis (Arlé, 1959) RJ NCB un [38,46] Insects 2021, 12, 268 32 of 39

Table 11. Cont.

Family/Species Brazil Distribution World Distribution Habitats Record Hypogastruridae Börner, 1906 Acherontides eleonorae Guano piles of hematophagous Palacios-Vargas and PR, SP NCB [53] Gnaspini-Netto, 1992 bats in caves Acherontides serrasapoensis Lima, Superﬁcial subterranean habitats MG NCB [54]* Stievano and Zeppelini, 2019 and cave Acherontiella colotipana ES Palacios-Vargas and Thibaud, 1985 Pal, Neo Soil [55] Acherontiella globulata Halophyte–psammophyte Thibaud and RJ Neo [37] Massoud, 1980 vegetation, sand dunes Austrogastrura marambaia Fernandes, Bellini and Mendonça, RJ NCB Halophyte–psammophyte [56] 2010 vegetation Halophyte–psammophyte Austrogastrura travassosi (Arlé, MS, RJ, PB Neo 1939) vegetation, litter, over rocks and [56–59] littoral sand Ceratophysella armata (Nicolet, Bor, Cos?, Neo un 1842) (RJ) [60] Ceratophysella bengtssoni (Ågren, RJ Bor, NCB Littoral sand [59] 1904) Ceratophysella engadinensis Leaf litter and soil of “campos de (Gisin PR, RJ Neo, Pal [61]* 1949) altitude” Ceratophysella nataliae Cipola, PR NCB Soil, semi-arid vegetation [62]* Bellini and Palacios-Vargas, 2017 Ceratophysella rogerarlei Soil, anthropized forest from Palacios- PI NCB [62]* Vargas, Bellini and Cipola, 2017 Atlantic forest Hypogastrura manubrialis (Tullberg, RJ Cos un [63] 1869) Hypogastrura rehi Börner, 1906 (SP) NCB un [36] Mesogastrura ojcoviensis (Stach, RJ Bor, Neo Littoral sand [59] 1919) Paraxenylla piloua Thibaud and RJ NCB Halophyte–psammophyte Weiner, 1997 vegetation [64] Paraxenylla sooretamensis Queiroz ES NCB and Deharveng, 2008 Forest litter [65] Paraxenylla zelliae sp.nov. AR NCB Oceanic islands sp. nov.* Schoetella celiae Fernandes and SP NCB Mendonça, 1998 Forest litter [66] Willemgastrura coeca Oliveira and AM, RO Amz, NCB Thibaud, 1988 Soil [67] Willemia brevispina Hüther, 1962 RJ, ES Ant, Neo, Pal Soil and littoral sand [55,59] Willemia insularum sp. nov FN, AR NCB Oceanic islands sp. nov.* Willemia zeppelinii D’Hese and PB NCB Thibaud, 2011 Coastal interstitial psammophilous [23] Leaf litter, forest soil, foredunes zone, halophyte-psammophyte Xenylla brasiliensis MG, RJ NCB da Gama, 1978 vegetation, dunes vegetation and [43,68]* supralittoral vegetation Xenylla capixaba Fernandes and ES NCB Mendonça, 2010 Leaf litter, sand dunes [42] Xenylla hodori Neves and AM Amz Mendonça, 2017 Leaf litter and soil [69]* Xenylla manuelae Queiroz and MG, ES, RJ NCB Mendonça, 2016 Soil leaf litter, altitude 2400 m [61]* Halophyte–psammophyte Xenylla maritima Tullberg, 1869 RJ Cos vegetation, sand dunes, ﬂooded [37] areas Xenylla nirae Gama and Oliveira, AM Amz 1994 Soil [70] Xenylla wandae Queiroz and RJ NCB Soil and leaf litter of altitude, Mendonça, 2016 2100 m [61]* Soil, halophyte– Xenylla welchi Folsom, 1916 ES, RJ Cos psammophytevegetation, [55,64] foredunes zone Beach sand, halophyte-psammophyte Xenylla yucatana Mills, 1938 FN, ES, RJ Pal, Neo, Aus vegetation, oceanic islands, dunes [3,42,43]* vegetation and supralittoral vegetation Isotogastruridae Thibaud and Najt, 1992 Isotogastrura mucrospatulata Palacios-Vargas, Lima and FN NCB Oceanic islands [34]* Zeppelini, 2013 Isotogastrura praiana halophyte-psammophyte Silveira, RJ NCB [71]* Mendonça and Da-Silva, 2014 vegetation and dunes vegetation Insects 2021, 12, 268 33 of 39

Table 11. Cont.

Family/Species Brazil Distribution World Distribution Habitats Record Neanuridae Börner, 1901 Aethiopella delamarei Arlé, 1960 MG, RJ, RN Neo Leaf litter [41,46,72] Halophyte–psammophyte Aethiopella littoralis Fernandes and vegetation, sand dunes, flooded Mendonça, 2002 RJ NCB Aethiopella ricardoi Paz, Bellini and RN NCB areas sandy soil surrounded by [73,74]* Queiroz, 2019 dead foliage near vegetation and lentic freshwater Anurida maritima Brazilian coast, PE, SP, Cos (Guérin-Méneville, 1836) RJ, ES, RN Surface of rock-pools on the coast [41,72,75–77] Arlesia albipes PA, PE, RJ, PB, PI, FN Neo Soil, leaf litter, sand dunes and (Folsom, 1927) forest, oceanic islands [38,43,78]* Arlesia arleana Mendonça and PE NCB Fernandes, 1999 Leaf litter [16] Arlesia intermedia Fernandes and RJ NCB Mendonça, 2004 Sand dunes, flooded areas [37] Arlesiella amazonica Arlé, 1966 AM Amz Leaf litter [72,79] Australonura gili Leaf litter and soil of “campos de Queiroz and RJ NCB [80]* Deharveng, 2014 altitude” Australanura neotropica Queiroz Leaf litter and soil of “campos de RJ NCB [80]* and Deharveng, 2014 altitude” Brasilimeria anura Arlé, 1939) RJ NCB Humus under dead leaf [81] Brasilimeria assu Queiroz and RJ NCB Leaf litter and soil of Brazilian Zeppelini, 2019 Páramos [82]* Brasilimeria wygodzinskyi (Arlé, RJ, MG NCB un 1943) [83,84] Cephalachorutes anneae Queiroz and RJ NCB Mendonça, 2016 Soil leaf litter, altitude 2500 m [61]* Ectonura snowdeni Queiroz and MG NCB Leaf litter and soil, altitude Deharveng, 2015 2500–2800 m [85]* Friesea arlei Massoud and Bellinger, MT Neo un 1963 [79] Friesea multiclavata Neves, AM NCB Forest leaf litter of Amazon Mendonça and Queiroz, 2019 Rainforest, [86]* Friesea boitata Queiroz and RJ Neo Mendonça, 2015 Soil and leaf litter 2582 m [87]* Friesea claviseta RJ Cos Halophyte–psammophyte Axelson, 1900 vegetation [64] Friesea cubensis Potapov and RJ Neo Littoral sand [59] Banasko, 1985 Friesea curupira Queiroz and RJ NCB Mendonça, 2015 Soil and leaf litter 2500 m [87]* Friesea josei Palacios- Vargas, 1986 RJ Neo Littoral sand [59] Friesea jurubatiba Beach sand and litter of “restinga” Silveira and RJ NCB [88]* Mendonça, 2018 soil Friesea magnicornis Denis, 1931 RJ Neo Halophyte–psammophyte vegetation [64] Friesea mirabilis RJ Neo Halophyte–psammophyte (Tullberg, 1871) vegetation [64] Friesea noronhaensis sp. nov. FN NCB Oceanic islands sp. nov.* Neo Halophyte– Friesea reducta Denis, 1931 RJ Neo psammophytevegetation, sand [37,64] dunes Friesea rochedoensis sp.nov. SPSP NCB Oceanic islands sp. nov.* Friesea sublimis Macnamara, 1921 ES Neo, Bor, Pal Soil [55] Furculanurida belemensis Arlé and PA Amz Rufino, 1976 Soil [83] Furculanurida boiunia Neves, AM Amz Forest leaf litter of Amazon Mendonça and Queiroz, 2019 rainforest [86]* Furculanurida goeldiana Arlé and PA Amz Rufino, 1976 Leaf litter [83] Furculanurida nessimiani Fernandes SP NCB and Mendonça, 2002 Forest litter [74]) Furculanurida tropicalia Queiroz ES NCB and Fernandes, 2011 Forest litter [89] Halachorutes schusteri Arlé, 1966 PA, RJ Amz, NCB Leaf litter [72,77] Handschinurida fluminensis (Arlé, RJ, SE NCB un [38,81] 1939) Handschinurida proxima (Arlé, 1939) RJ, SE, SP NCB un [16,38,81] Handschinurida rauli Queiroz and MG NCB Mendonça, 2014 Soil and leaf litter, altitude 2700 m [90]* Hylaeanura infima MT, AM, PA, RJ Neo Soil, leaf litter, sand dunes, flooded (Arlé, 1959) areas [37,38,46,79] Hylaenura mendoncae Zeppelini and Areas of iron ore mining; leaf litter MG NCB of semi-deciduous forest [91] Palacios- Vargas, 2013 fragments Itanura brasiliensis (Arlé, 1960) RJ, MG NCB [92] Kenyura delicata Arlé, 1966 AM Amz Leaf litter [72,79] Kenyura porcula (Arlé, 1959) RJ NCB [46] Kenyura xinguensis Arlé, 1966 MT NCB Leaf litter [72,79] Insects 2021, 12, 268 34 of 39

Table 11. Cont.

Family/Species Brazil Distribution World Distribution Habitats Record Micranurida fluminensis Fernandes RJ NCB and Mendonça, 2004 Sand dunes, flooded areas [37]) Neotropiella arlei Primary and secondary forest soil Najt, Thibaud and AM Neo [93] Weiner, 1990 and litter Neotropiella barbatae Queiroz, Soil and leaf-litter of “campos de RJ NCB [94]* Silveira and Mendonça, 2013 altitude” 2400m Neotropiella carli (Denis, 1924) (AM, AP, PA) Neo Leaf litter [38] Neotropiella denisi (Arlé, 1939) RJ, SE, MT NCB Soil, litter on roots [38,72,81] Neotropiella digitomucronata Primary and secondary forest soil AM Neo [93] Thibaud and Massoud, 1983 and litter Neotropiella insularis Queiroz, RJ NCB Silveira and Mendonça, 2013 Leaf litter of Atlantic rainforest [94]* Neotropiella macunaimae Queiroz, MG NCB Silveira and Mendonça, 2013 Soil and leaf litter, altitude 2700 m [94]* Neotropiella meridionalis (Arlé, 1939) SE, MT, RJ, MG, PA, AM Neo Leaf litter, rotten woods [38,72,79,81,93] Neotropiella minima Thibaud and AM Amz Oliveira, 2011 Floodable rain forest, clay soil [95] Neotropiella plurichaetosa Thibaud AM Amz and Oliveira, 2011 Floodable rain forest, clay soil [95] Neotropiella quinqueoculata (Denis, RJ, AM, AP, MT, PA, RJ Bor, Neo un 1931) [35,79] Neotropiella silvestrii (Denis, 1929) AM Neo [95] Neotropiella vanderdrifti Primary and secondary forest soil Massoud, AM Neo [93] 1963 and litter Paleonura brasiliensis (Arlé, 1959) MG (RJ) NCB Leaf litter [46] Paleonura nuda Cassagnau and Primary and secondary forest soil AM Amz [96] Oliveira, 1990 and litter Pronura amazonica Cassagnau and AM Amz Leaf litter [96] Oliveira, 1990 Pseudachorutes bifasciatus Oliveira AM Amz and Deharveng, 1994 Soil [97] Pseudachorutes difficilis Denis, 1931 RJ, MG, ES Neo Sand dunes, flooded areas, soil and leaf litter of highlands grasslands [37,43]* Pseudachorutes gilvus Oliveira and AM Amz Deharveng, 1994 Soil [97] Pseudachorutes herberti Arlé and AM NCB Rufino, 1976 Leaf litter [83] Pseudachorutes massoudi Arlé, 1966 AM Amz Leaf litter [72,79] Pseudachorutes parvulus PB Cos un Börner,1901 [98] Pseudachorutes solaris Silveira and RJ NCB Mendonça, 2018 Sand beach and litter of “restinga” [88]* Pseudanurida sawayana Schuster, On sand and seaweed on the rocky 1965 PE, SP, (RJ) Bor, Neo, Pal coast [76] Leaf litter and soil between roots Tijucameria gabrieli Mendonça and RJ NCB on montane vegetation of the Silveira, 2012 [99]* Atlantic rainforest Tijucameria mame Mendonça and RJ NCB Fernandes, 2005 Forest litter [100] Odontellidae Massoud, 1967 Stachia folsomi (Arlé, 1968) PA, RJ Amz, Pal Soil, dunes vegetation [43,101]* Onychiuridae Lubbock, 1867 Agraphorura fernandae (Oliveira PA Amz and Thibaud, 1992) Leaf litter [102] Agraphorura mariapetrae (Thibaud, RJ Neo 1993) Flooded areas [37] Onychiurus cunhai Arlé, 1970 PA, AM Neo Soil [58,103] Protaphorura cryptopyga (Denis, RJ un 1931) Neo, Pal [35] Tullbergiidae Bagnall, 1935 Soil, halophyte– Fissuraphorura cubanica Rusek, 1991 RJ, ES Neo [59,63] psammophytevegetation Mesaphorura amazonica Litter, halophyte–psammophyte Oliveira AM, RJ, ES Amz, NCB [37,55,102] and Thibaud, 1992 vegetation, sand dunes, soil Mesaphorura iowensis (Mills, 1932) SP Cos Soil and litter [104] Mesaphorura maricaensis Fernandes RJ NCB and Mendonça, 2004 Sand dunes [37] Primary and secondary forest soil and litter; Mesaphorura yosiii (Rusek, 1967) AM, SP, RJ Cos halophyte–psammophyte [59,102,104] vegetation, sand dunes and flooded areas, soil, litter Tullbergia minensis Arlé, 1959 MG NCB Litter [46] Insects 2021, 12, 268 35 of 39

4. Discussion The ﬁrst Collembola species registered in Brazil was the Seira musarum Ridley, 1890 (Entomobryomorpha), whose type locality is Fernando de Noronha. However, after this pioneering study, the only works with Collembola in Brazilian oceanic islands were the description of Isotogastrura mucrospatulata Palacios, Lima and Zeppelini 2012 [36] and the species survey done by Lima and Zeppelini, 2015 [3] both in Fernando de Noronha. Our results show that Poduromorpha fauna of the oceanic islands showed a high percentage of species known only from their type localities and with a great potential for endemisms. All of the Poduromorpha species found in the São Pedro and São Paulo archipelago and Atoll das Rocas are undescribed and with consistent morphological differences in relation to the species found on the continent. Fernando de Noronha, the only Brazilian oceanic island where tourist visitation is allowed, one endemic (I. mucruspatulata), four with wide continental distribution and two new species described here. These results may be useful in the management and environmental monitoring programs of oceanic islands in Brazil. Although the increase in the number of Brazilian Poduromorpha species over the past decade has been relevant (about 35% since [2]) when we consider the variety of Brazilian continental biomes and the complex physiognomy of these areas, current knowledge of the Poduromorpha fauna can still be considered scarce and far from providing a reliable overview of its biogeographical panorama.

Author Contributions: Conceptualization, all authors; methodology, E.C.A.d.L.; formal analysis, all authors; investigation, E.C.A.d.L. and D.Z.; resources, all authors; data curation, D.Z.; writing— original draft preparation, E.C.A.d.L. and T.C.d.S.; writing—review and editing, E.C.A.d.L. and G.C.Q.; supervision, M.C.d.M. and D.Z.; project administration, E.C.A.d.L.; funding acquisition, all authors. All authors have read and agreed to the published version of the manuscript. Funding: E.C.A.d. Lima is granted by CNPq (Protax, Process 440521/2015-7), M.C.d. Mendonça is granted by CNPq (Process 307644/2015-4), D Zeppelini is granted by CNPq (Process 301803/2012-9). Institutional Review Board Statement: The study was conducted according to the guidelines of the Declaration of Helsinki and approved by the Institutional Review Board of Chico Mendes Institute for Biodiversity Conservation (ICMBio) (SISBIO 32469). Informed Consent Statement: Not applicable. Data Availability Statement: All the specimens collected and used in this research are deposited in the Coleção de Referência de Fauna de Solo at the State University of Paraiba (CRFS-UEPB) under tumble number as cited in the materials and methods. Acknowledgments: The ICMBio provided the permits for collection; Marinha do Brasil provided the transportation to St Peter and St Paul rocks, the preparatory training, and support during the scientific expeditions; Aíla Soares Ferreira and Diego Dias da Silva gave support during the fieldwork in Fernando de Noronha; Luís Carlos Stievano helped in the ordenation of the samples and assistanced in all laboratory work; anonymous reviwers greatly improved the manuscript. A special thanks for the crew members of the boats Borandá and Alfa, for passages to the most incredible places I have yet had the opportunity to know: the Oceanic Islands. Conflicts of Interest: The authors declare no conflict of interest.

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