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Zhang, Feng, Bellini, Bruno C., Soto-Adames, Felipe N. (2019): New insights into the systematics of Entomobryoidea (Collembola: Entomobryomorpha): first instar chaetotaxy, homology and classification. Zoological Systematics 44 (4): 249-278, DOI: 10.11865/zs.201926, URL: http://dx.doi.org/10.5281/zenodo.4421087

Plutomurus pichkhaiai Barjadze, Jordana & Soto-Adames sp. nov. Figs 23–26 Type locality. GEORGIA, Samegrelo– – Zemo Svaneti, Chkhorotsku, near Garakha, Garakha Cave, 42°31'47.22"N, 42°10'39.18"E, 203 m alt. Type material. Holotype, female on slide: dark zone, 13.ii.2014, leg. Sh. Barjadze (code GeoGar20140213– 02). Paratypes (same as holotype): 2 females on slides (code GEOShu20140213–01 and 03). Two specimens mounted on SEM stubs (GeoGar20140213–04 and 05). Three specimens in ethanol (GeoGar20140213–06, 07 and 08). Paratypes: two females on slide, dark zone, 30.iv.2017, leg. G. Nebieridze (GeoGar20170430–01 and 02); one female on slide, twilight zone, 30.iv.2017, leg. G. Nebieridze (GeoGar20170430–03). Two specimens (FSCA 108– 109, males) same date as holotype but 30.iv.2017, leg. G. Nebieridze. The material from 13.ii.2014 was previously mentioned as “ Plutomurus sp. 4” in Barjadze et al. (2015) and as “ Plutomurus sp. 1” in Barjadze et al. (2016). Additional material. GEORGIA, Samegrelo–Zemo Svaneti, Chkhorotsku, Letsurtsume, Letsurtsume Cave, 42°32'21.4"N, 42°06'47.3"E, 183 m alt. Two specimens (FSCA 110, female and FSCA 111 male) on slides, dark zone, 29.iv.2017, leg. Sh. Barjadze and two specimens (GeoLet20170429–01, male) and (GeoLet20170429–02, female) on slides, dark zone, 29.iv.2017, leg. Sh. Barjadze. Holotype and paratypes 01, 03–08 deposited at MZNA; paratypes: GeoGar20170430–01, 02, 03 at IZISU, and N108–109 at FSCA. Additional materials (GeoLet20170429–01 and 02) at IZISU, and N110–111 at FSCA. Description. Body length up to 3.76 mm excluding antennae. Color pattern. Habitus and distribution of grey pigment as in Fig. 23. Scale distribution. As in P. shurubumuensis sp. nov. Head. Ratio of body to antennae length up to 0.85. Number of eyes unclear, observations under compound microscope show 3–5 eyes, but field where eyes sit often distorted. Head dorsally with 1 unpaired, and 6 paired Mc as in P. shurubumuensis sp. nov. Prelabral chaetae 3+3, labral chaetae and papillae typical for genus. Maxilla as in P. shurubumuensis sp. nov., but lamella N5 sometimes appearing Y-shaped (Fig. 24) Proximal chaetae of labium 16–17. Basomedial field with 22–23 smooth chaetae; basolateral field with 5 chaetae. Body. Dorsal Mc formula 4,2//3,3,4,3,4 (Figs 25–26). Abd. IV with 7 differentiated chaetae, chaeta #7 always a large Mc, chaetae N2 and N3 always developed into a small Mc (Fig. 26), other chaetae always small, mesochaeta-like as in P. shurubumuensis sp. nov. Legs. Hind legs with well-developed trochanteral (16–30 chaetae) and femoral organs (25–36 chaetae) (n=7). Tibiotarsal outstanding spine-like chaetae formula as 002. Tenent hair acuminated, as in P. shurubumensis sp. nov. Ratio hind unguis: unguiculus: tenent hair as 1.50–3.00: 1.50–2.28: 1 (n=7). Inner edge of unguis on all legs with 2 unpaired teeth, proximal tooth minute, distal tooth evidently larger; lateral outer teeth 0.38–0.83 times length of inner edge of uguis (n=7). Unguiculus lanceolate and tapered, with 0–7 inner teeth. Collophore. Chaetotaxy as in P. shurubumuensis sp. nov. Furcula. Ratio manubrium: dens: mucro as 3.52–5.00: 7.17–8.17: 1 (n=6, including holotype). Outer margin of dens with 3–5 acuminate spine–like macrochaetae. Inner edge of dens with a variable number of basal spines; row of spines on distal segment of dens extending between 24–28% (3 out of 6 individuals=26%) of dens total length. Dental spines formula variable as 5–14 II–IV /8–13 II–IV. Mucronal teeth formula 202. Variation. The number of teeth on inner lamellae of the unguiculus varies from 0 to 7. The number of dental spines is also variable. Ratio of hind unguis: unguiculus: tenent hair, and ratio of manubrium: dens: mucro are also variable. Discussion. Plutomurus pichkhaiai sp. nov. is the only eyed member of the genus with 3+3 prelabral chaetae, 2 spines on the inner edge of the hind legs, and 3 posterior Mc on Abd. IV. The new species is most similar to P. shurubumuensis sp. nov., from which it differs in the number of posterior Mc on Abd. IV, in mtDNA sequence and cave formation as described above for P. shurubumuensis sp. nov. From P. kelasuricus the new species differs in having 2 inner spine–like chaetae on the inner edge of the hind tibiotarsus (only 1 in P. kelasuricus); in addition, P. pichkaiai sp. nov. was collected in Garakha and Letsutsume caves, part of the Migaria karst massif formation, whereas P. kelasuricus was collected 100 km to the northwest, in Kelasuri Cave, which is part of the Tsebelda karst massif formation. Etymology. The epithet honours Mr. Igor Pichkhaia, who has worked on the popularization of local caves in Chkhorotsku district during several decades. Ecology. The presence of body and eye patch pigmentation suggest this is a troglophilous species.

Plutomurus shurubumuensis Barjadze, Jordana & Soto-Adames sp. nov. Figs 4���22 Type locality. GEORGIA, Samegrelo��� ��� Zemo Svaneti region, Chkhorotsku district, 3 km from Mukhuri, Migaria karst massif, Shurubumu Cave, 4239'5.71"N, 4213'44.58"E, 464 m alt. Type material. Holotype, female on slide: dark zone, 03.x.2014, leg. Sh. Barjadze (code GEOShu20141003��� 04). Paratypes (same data as holotype): four females on slides (code GEOShu20141003���01, 02, 03 and 05). Three specimens mounted on SEM stubs (GEOShu20141003���06, 07 and 08). Seven specimens in ethyl alcohol (GEOShu20141003���09���15); Paratypes (same data as holotype but 28.ix.2017) one male (GEOShu20170428���01) and one female (GEOShu20170428���02) on slides, leg. Sh. Barjadze; three additional females with same date as holotype but 28.ix.2017, leg. Sh. Barjadze. Holotype and paratypes 01, 02, 06, 07 and 08 deposited at MZNA; paratypes GEOShu20141003���03 and 05, GEOShu2017040328���01 and 02 deposited at IZISU; 3 female paratypes deposited at FSCA (N105���107). Description. Body length up to 3.36 mm, excluding antennae and furcula. Color pattern. Habitus and grey pigment distribution as in P. pichkhaiai sp. nov. (Fig. 23). Scale distribution. Dorsally on Ant. I���II, head, body, all legs segments, both faces of collophore and ventral face of furcula. Postlabial region of head with few or no scales. Head. Ratio body length to antennae length up to 1.09. Number of eyes unclear, apparently 6, but all cornea reduced and actual number difficult to determine in most specimens; under light microscope eye number appears to vary from 3 to 6, whereas the single individual evaluated using SEM clearly shows 6 eyes (Fig. 4). Head dorsally (Fig. 7) with one unpaired (A 0), and 6 (7) paired Mc: 2 anterior (A 2, A 3), 2 interocular (Ps 2, Ps 3) and 2(3) postocular ((Pa 2), Pa 3, Pa 5) (supplementary Mc represented by dotted line in Fig. 7). Prelabral and labral chaetae smooth (Fig. 11): prelabral chaetae 6 (3+3); labrum with 554 papillate chaetae as typical for genus; distal margin of labrum with 4 elongate, thin-walled, curved papillae. Outer maxillary lobe trifurcate (Fig. 12), basal chaetae shorter than apical process; sublobal plate with 4 processes each one resting on a short papilla. Sclerotized head of maxilla (Fig. 13) with 3 large and 1 small teeth; maxillary lamella 1 (Fig. 6) evidently longer and more slender than others, with a small posterior basal brush and better developed anterior medial brush, ventral margin with long cilia; lamella 2 (Fig. 6), and 4 (not shown) wide, densely covered with denticles and hooks; lamella 3 (Fig. 6) short, somewhat ellipsoidal, with cilia regularly distributed along its entire margin; lamella 5 (Fig. 14), sinuous but variable in shape, basal beard absent (i.e., prostheca not developed); lamella 6 narrow, comb-shaped, with short processes (Fig. 15). Labial papilla E with 6 guard processes; lateral modified process thin-walled, tapering and surpassing tip of papilla (Fig. 16 inset). Labium with 16���17 proximal chaetae (Fig. 16). Basomedial field (i.e., labial triangle) without scales, with 22���23 smooth chaetae; basolateral field with 5 chaetae (A3���A5 and L1���L2), chaetae on anterior row slightly longer than those on posterior row. Body. Dorsal bothriotrichal formula 2,1//0,0,1,2,0 (Figs 8���9). Dorsal Mc formula 4,2//3,3,4,1(2),4 (Figs 8���10). Thorax macrochaetotaxy as in Fig. 8: Th. II with 2 medial and 2 posterior Mc; Th. III with 2 posterior Mc. Abdominal macrochaetotaxy as in Figs 9���10: Abd. I��� ���II each with 3 posterior Mc; Abd. III with 2 anterior and 2 posterior Mc as typical for genus; Abd. IV with 7 enlarged chaetae (labeled 1���7 in Figs 9���10), chaeta N7 always largest Mc, chaeta N2 a small Mc in 1 out of 6 individuals examined, all other differentiated posterior chaetae clearly enlarged, but sockets always small, mesochaeta-like; Abd. V with 4 posterior Mc, 1 additional lateral Mc always present, but often hidden by slide mounting induced deformation of cuticle, and segment appears as having 4 Mc. Legs. Hind legs with well-developed trochanteral (18���35 chaetae) and femoral organs (20���44 chaetae) (n=9) (Fig. 17). Posterior face of hind tibiotarsus with 2 outstanding pointed spine-like chaetae, one basal and one distal, resulting in a 002 tibiotarsal spine formula (Fig. 20). Tenent hair acuminated, thinner and shorter than flanking chaetae, and inserted in a well-developed depression (Fig. 18). Ratio hind unguis: unguiculus: tenent hair as 1.54��� 3.55: 0.92���2.48: 1 (n=11). Inner edge of unguis on all legs with one minute proximal unpaired tooth (a in Figs 5 and 18), and 1 larger distal unpaired tooth (b in Figs 5 and 18). Unguis with lateral teeth 0.42���0.86 as long as length of inner edge (n=9), reaching proximal minute inner tooth. Unguiculus lanceolate, tapered, with 2 internal lamellae bearing 0���6 teeth (Fig. 18). Collophore. Anterior, posterior and distal faces with 27, 45 and ca. 65 smooth chaetae, respectively. Tenaculum. Corpus with 1 smooth chaeta; rami with 4 + 4 teeth (Fig 19). Furcula. Ratio manubrium: dens: mucro as 3.83���6.00: 4.80���8.75: 1 (n=8, including holotype). Outer margin of basal segment of dens with 3(4) apically acuminate macrochaetae, distal macrochaeta largest, proximal shortest (Fig. 22). Inner edge of dens basally with well differentiated spine-like chaetae; spines on basal segment of dens forming 2���3 short and poorly organized rows, upper row usually formed by 2 large, well-differentiated spines; spines on distal segment forming a single row extending distally between 35���43% of dens length; 3 proximal spines always small, terminal spine always largest in row; with long spines intercalated between short spines (Fig. 22), but always with at least 2 (sometimes 3) distinctly larger distal spines; total dental spines formula variable as 9���13 II���IV /10���13 III���IV. Mucro with 2 proximal and 2 distal teeth (202 formula): proximal teeth fused throughout most of their length, separated only at tip. (Fig. 21). Variation. The actual number of eyes is difficult to ascertain using light microscope preparations. The dorsal cephalic and Abd. IV chaetotaxies are variable: one individual has three posterior Mc on the head instead of the two normal Mc; several individuals have a distinctly enlarged dorsal posterior chaeta N2 on Abd. IV. The number of teeth on the inner lamellae of the unguiculus varies from 0 to 6. The total number of dental spines also varies as reported above. Ratio of unguis III: unguiculus III: tenent hair and ratio of manubrium: dens: mucro are variable characters. Discussion. Among members of the genus Plutomurus, only P. shurubumuensis sp. nov. has eyes, 3+3 prelabral chaetae, 2 spines on the inner edge of metathoracic legs, and 1 (sometimes 2) posterior Mc on Abd. IV. The new species is most similar to P. pichkhaiai sp. nov., and P. kelasuricus. The new species differs from P. pichkhaiai sp. nov., in the number of posterior Mc on Abd. IV (1���2 in P. shurubumuensis sp. nov., 3 in P. pichkhaiai sp. nov.) a p-distance of at least 13.7% in the 3��� half of the COI mitochondrial gene, which results in reciprocally exclusive monophyletic sister groups in a Bayesian analysis (cf. Barjadze et al. 2016), and in that they occupy different cave formations (see Materials & Methods) isolated by a fault. From P. kelasuricus the new species differs in having 2 inner spine���like chaetae on the inner edge of the hind tibiotarsus (only 1 in P. kelasuricus) and in occupying different, isolated cave formations separated by a geographic distance of nearly a 100 km (Fig. 1). Plutomurus shurubumuensis sp. nov. is also very similar to P. revazi, which has a capitate tenent hair (acuminate in the new species), and 5 (medial and posterior), 3 Mc on Th. II and Abd. V, respectively (4, 4 Mc in the new species). All other seven species of Plutomurus with 3+3 prelabral chaetae (P. ehimensis Yosii, 1956, P. eristoi, P. gul (Yosii, 1966), P. grahami, P. iwatensis Yoshii, 1991, P. kawasawai Yosii, 1956, and P. ortobalaganensis) are blind and easily distinguished from P. shurubumuensis sp. nov. Etymology. The species is named after the cave in which it was collected. Ecology. The body and eye patch pigmentation, together with the thick, toothed unguis suggest this is a troglophilous species. The small number and size of the inner ungual teeth and presence of an acuminate tenent hair place it in the early stages of troglomorphy., Published as part of Barjadze, Shalva, Jordana, Rafael, Baquero, Enrique, Giordano, Rosanna & Soto-Adames, Felipe N., 2018, Two new species of Plutomurus Yosii (Collembola, Tomoceridae) from the Caucasus, pp. 29-40 in Zootaxa 4526 (1) on pages 32-36, DOI: 10.11646/zootaxa.4526.1.2, http://zenodo.org/record/3771190, {"references":["Barjadze, Sh., Baquero, E., Soto-Adames, F., Giordano, R. & Jordana, R. (2016) New diagnosis for species of Plutomurus Yosii (Collembola, Tomoceridae), with descriptions of two new species from Georgian caves. Zootaxa, 4126 (1), 77 - 96. https: // doi. org / 10.11646 / zootaxa. 4126.1.3","Yosii, R. (1956) Monographie zur Hohlencollembolen Japans. Contribution from the Biological Laboratory Kyoto University, 3, 1 - 109.","Yosii, R. (1966) Results of the Speleological Survey in South Korea 1966. IV. Cave Collembola of South Korea. Bulletin of the National Science Museum, 4, 541 - 561.","Yoshii, R. (1991) About the proserpinae group of Hypogastrura (Collembola) in the Caves of Pref. Iwate. Annals of the Speleological Research Institute of Japan (Iwaizumi), 9, 3 - 10."]}

Key to the species of the genus Plutomurus with 3+3 prelabral chaetae Most characters used to diagnose Plutomurus species are ambiguous and open to interpretation. Only the number of prelabral chaeta is easy to see, discrete, and with insignificant levels of intraspecific variation. As discussed above, under light microscopy it is often difficult to determine the actual number of eyes present, although at least the 2���3 largest eyes are almost always visible. Hence, the key refers only to eyes present or absent. 1 Eyes present (usually 3���6)............................................................................. 2 - Eyes absent......................................................................................... 5 2 Hind tibiotarsus with 1 posterior spine.......................................................... P. kelasuricus - Hind tibiotarsus with 2 posterior spines................................................................... 3 3 Tenent hair capitate; Th. II and Abd. V with 5 (medial & posterior) and 3 (posterior) Mc, respectively............ P. revazi - Tenent hair acuminate; Th. II and Abd. V each with 4 Mc...................................................... 4 4 Abd. IV with 1 large lateral and sometimes a second smaller medial Mc (Figs 9���10).......... P. shuburumensis sp. nov. - Abd. IV with 3 Mc: 1 large lateral and 2 smaller medial Mc (Fig. 26)......................... ��� P. pichkhaiai sp. nov. 5 Tenent hair capitate............................................................................ P. grahami - Tenent hair acuminate.................................................................................. 6 6 Mucro with 1���3 intermediate teeth....................................................................... 7 - Mucro without intermediate teeth........................................................................ 8 7 Mucro with 1 proximal tooth................................................................... P. ehimensis - Mucro with 2 proximal teeth......................................................................... P. gul 8 Tenaculum with 2 chaetae; apical and subapical teeth of mucro truncate................................. P. iwatensis - Tenaculum with 1 chaeta; apical and subapical teeth of mucro rounded........................................... 9 9 Inner edge of hind unguis toothless; unguiculus basally swollen and medially excavated...................... P. eristoi - Hind unguis with 1���3 inner teeth; unguiculus narrower at base than near middle................................... 10 10 Hind tibiotarsus with 1 inner spine; unguiculus inner teeth present............................. ��� P. orthobalaganesis - Hind tibiotarsus with 2 inner spines; unguiculus inner teeth absent.................................... P. kawasawai, Published as part of Barjadze, Shalva, Jordana, Rafael, Baquero, Enrique, Giordano, Rosanna & Soto-Adames, Felipe N., 2018, Two new species of Plutomurus Yosii (Collembola, Tomoceridae) from the Caucasus, pp. 29-40 in Zootaxa 4526 (1) on page 38, DOI: 10.11646/zootaxa.4526.1.2, http://zenodo.org/record/3771190

Trogolaphysa grassei (Delamare Debouteville, 1952) n. comb. Paronella grassei Delamare Debouteville, 1952: 241, 242, figs 4-7 (Le Banco, Ivory Coast). Paronella grassei Delamare Debouteville – 1951b: 63, fig. 16, nomen nudum. Dicranocentruga grassei – Mitra 2002c: 114. TYPE LOCALITY. — Le Banco, Ivory Coast. DISTRIBUTION. — Ivory Coast. REMARKS Delamare Debouteville (1951b) first published a plate showing diagnostic characters for this species and included the combination Paronella sp. in the caption to the figure. However, this publication did not include a description of the species. Delamare Debouteville (1952) published the 1951b plate and caption again, but this time he included a formal description. Following the ICZN code, the absence of a description and diagnosis renders the 1951b combination a nomen nudum and the correct authority citation is P. grassei Delamare Debouteville, 1952. This species is placed in Trogolaphysa based on the absence of manubrial spines and on the shape of the mucro, which is typical for Trogolaphysa species with 3-teethed mucro. This species has a white head and light blue pigment on legs and body flanks; tenent hair relatively short and weakly spatulate (Fig. 5A); unguis with three inner teeth; unguiculus lanceolate; dens with two rows of spines; and mucro with 3 teeth. This is the only African species with a three-toothed mucro., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 183, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["DELAMARE DEBOUTTEVILLE C. 1952. - Trois nouvelles especes de Collemboles de la Cote d'Ivoire. Revue francaise d'Entomologie 19: 240 - 243.","DELAMARE DEBOUTTEVILLE C. 1951 b. - Microfaune du sol des pays temperes et tropicaux. Actualites scientifiques et industrielles 1160: 1 - 360.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116."]}

Genus Trogolaphysa Mills, 1938 sensu Soto-Adames (2015) Trogolaphysa Mills, 1938: 184. TYPE SPECIES. — Trogolaphysa maya Mills, 1938: 184, by original designation. TYPE LOCALITY. — Yucatan, Mexico. DIAGNOSIS. — Hyaline denticulated scales covering Ant. 1-2, head, body, and ventral face of furcula; 4th antennal segment not subdivided; eyes 0-8; head microchaeta Ps 3 absent; labial chaetae L1 and L2 subequal; inner dorsal chaetotaxy of metathorax and 1 st abdominal segment reduced to 4 and 3 chaetae, respectively; abdominal segments 2-4 with 2, 3, 3 bothriotricha; manubrium without spines; dens with 1-2 rows of spines; mucro square or rectangular but relatively short, with 3-5 teeth. REMARKS Published descriptions of species listed below as belonging to Trogolaphysa (except for D. wahlgreni and the new species) do not refer to most characters listed in the diagnosis of the genus. For this study, we assign to Trogolaphysa all forms originally placed in the genus Paronella Schött, 1893 that, based on available descriptions, have a square or short rectangular mucro bearing 3-5 teeth, and lack manubrial spines and a subdivision on the 4 th antennomere. This limited combination of characters is not unique to Trogolaphysa, as it would also fit some members of the genus Lepidonella Yosii, 1960. Trogolaphysa and Lepidonella are easily distinguished based on chaetotaxy, and to a large extent, on mucro shape. Most species of Trogolaphysa (at least the Neotropical forms) have the typical relatively short (or even shorter) mucro shown in Fig. 3C, whereas in Lepidonella (and more so in Troglopedetes Absolon, 1907 / Cyphoderopsis Carpenter, 1917) the mucro tends to be appreciably longer. A few species of Lepidonella do have short mucrones, but most of these are shaped very differently from those in Trogolaphysa (Fig. 3C; Soto-Adames & Bellini 2015: figs 28, 29). Only Lepidonella ceylonica (Yoshi, 1966), and L. nigrofasciata (Handschin, 1928) have Trogolaphysa -like short mucrones, and both species are found in South Asia. Lepidonella has never been reported from the African mainland; only L. flava (Carpenter, 1916; from the Seychelles) has been reported from a region near Africa. The chaetotaxy of most species listed below is not known but the mucro is always illustrated. Although we use mucro shape as the main character to assign species to Trogolaphysa, other characters, when available, are used in support of our determination. Thus, the description of Lepidonella flava is not sufficiently detailed to assigned it to Lepidonella or Trogolaphysa, but we have retained it in Lepidonella based on one aspect of the chaetotaxy. We present our comments on L. flava following the identification key to African Trogolaphysa. Trogolaphysa vivieni (Barra, 1969) and T. monomaculata have distinctive mucrones (especially T. viviani), which are atypical for the genus, but Soto-Adames & Bellini (2015) transferred them to Trogolaphysa on account of the number of bothriotricha on the 4th abdominal segment and absence of head chaeta Ps 3. Justification for placing other species in Trogolaphysa is given in the ‘Remarks’ to each form. ANNOTATED CHECKLIST OF AFRICAN TROGOLAPHYSA Thibaud’s (2013) report on African springtails lists 17 species and three subspecies in the genera Paronella, Microparonella Carpenter, 1916 and Trichorypha Schött, 1893 that may belong in Trogolaphysa (Table 1). We evaluated published descriptions of the species listed in Thibaud (2013) in light of current generic delimitation and the distinguishing criteria listed above, and our conclusions are presented below (Table 1). Following Deharveng & Bedos (1995) Microparonella caerulea Carpenter, 1916 belongs in Troglopedetes. Mitra (2002a) studied a syntype of M. caerulea Carpenter, 1916 and suggested that Microparonella was a valid genus. However, the specimen Mitra examined was missing the three distal antennal segments and he could not be aware of the diagnostic character placing the species in Troglopedetes (see Remarks below under L. flava). Paronella villiersi Delamare Debouteville & Pauliani, 1947 has been cited close to 10 times in the last 65 years, but the species was never described or illustrated and therefore the combination is a nomen nudum. Paronella penicillata obscura Delamare Debouteville, 1951, from Ivory Coast was never diagnosed and is also a nomen nudum. Paronella eburnea Delamare Debouteville & Paulian, 1947 is a nomen nudum, which may have referred to a species from Ivory Coast later described as Aphysa eburnea Delamare Debouteville, 1951. The shape of the mucro and length of antennae suggest that A. eburnea belongs in Callyntrura Börner, 1906 or at least in Callyntrurini Mitra, 1993. The generic status of Paronella atrofasciata (Schött, 1893) is unclear. Thibaud & Najt (1988) indicated that type material of P. atrofasciata they examined fit the diagnosis of Paronella in having 6+ 6 eyes, relatively short antennae and manubrial spines. However, Mitra (2002b) reported syntypes as having 8 + 8 eyes, antennae longer than body and manubrial spines absent. The absence of manubrial spines excludes the specimens Mitra examined from Paronella. The additional absence of extra-ocular structure (eos) in combination with other characters lead Mitra to resurrect the genus Trichorypha. The chaetotaxy for Abd. 2-4 illustrated by Mitra (2002b) for T. atrofasciata is unlike that seen in other Paronellini. A clarification of the status of this species will require examination of the chaetotaxy of the complete type series or fresh material from Cameroon. Paronella fusca Schött, 1893 (Type species of Paronella), P. montana Delamare Debouteville, 1952 and P. purpurea Barra, 1969 carry manubrial spines and belong to Paronella s. str. (Thibaud & Najt 1988). , The remaining 12 species of Paronella reported from Africa may be tentatively assigned to Trogolaphysa. The annotated checklist and identification key presented below are based on our evaluation of published descriptions and records. In the annotations we discuss some important papers concerning the taxonomy of each species, but it was not our intention to list all citations for all species. Synonymies, complete up to around 2000, for all forms are available from www. collembola.org (Bellinger et al. 1996 -2018) under links to ‘Publications’ and ‘ Collembola species catalogue’, Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on pages 181-182, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["MILLS H. B. 1938. - Collembola from Yucatan Caves. Carnegie Institution of Washington Publication 491: 183 - 190.","SCHOTT H. 1893. - Beitrage zur Kenntnis der Insektenfauna von Kameroon. I. Collembola. Bihang till Kongliga Svenska Vetenskaps- Akademien Handligar 19: 1 - 28.","CARPENTER G. H. 1917. - Collembola, zoological results of the Abor expedition, 1911 - 12. Records of the Indian Museum 8: 561 - 568.","HANDSCHIN E. 1928. - Collemboles aus Java, nebst einem beitrag zu einer monographie der gattung Crematocephalus Schtt. Treubia 10: 245 - 270.","CARPENTER G. H. 1916. - The Apterygota of the Seychelles. Proceedings of the Royal Irish Academy 33: 1 - 108.","BARRA J. - A. 1969. - Collemboles du Gabon. Revue Biologia Gabonica 5: 189 - 216.","THIBAUD J. - M. 2013. - Essay on the state of knowledge of collembolan diversity in the African-Malgasy Region. Russian Entomological Journal 22: 233 - 248.","DEHARVENG L. & BEDOS A. 1995. - Lepidonella lecongkieti n. sp., premiere Collembole cavernicole de Vietnam. Bulletin de la Societe entomologique de France 100: 21 - 24.","MITRA S. K. 2002 a. - Status of Microparonella Carpenter, 1916 (Collembola: Entomobriyidae: Paronellinae). Records of Zoological Survey of India 100: 67 - 72.","BORNER C. 1906. - Das System der Collembolen nebst Beschreibung neuer Collembolen des Hamburger Naturhistorischen Museums. Mitteilungen aus dem Naturhistorischen Museum 23: 147 - 188.","THIBAUD J. - M. & NAJT J. 1988. - Collemboles (Insecta) de l'Equateur IV. Paronellidae avec revision de quarter genres. Bulletin du Museum national d'Histoire naturelle 4: 719 - 730.","MITRA S. K. 2002 b. - Resuscitation of Trichorypha Schott, 1893 (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 99 - 104.","BORNER C. 1903. - Uber neue Altweltliche Collembolen, nebst Bermerkungen zur Systematik der Isotominen und Entomobryinen. Gesellschaft naturforschender Freunde: 129 - 182.","MARLIER G. 1945. - Collemboles du Congo belge. Revue de Zoologie et de Botanique africaines 38: 252 - 264.","SCHOTT H. 1903. - Uber zwei Gattungen der Apterygoten Insekten. Redogorelse for Allamanna Larvorken I Linkoping. Esko och Vadstena. Underlarsaret: 3 - 16.","SCHOTT H. 1927. - Kamerunische Collembola. Medd. Linkopings Hogre Allmanna Laroverk redogorelse: 1 - 39.","BELLINGER P. F., CHRISTIANSEN K. A. & JANSSENS F. 1996 - 2017. - Checklist of the Collembola of the World. http: // www. collembola. org. Last accessed 3 February 2018."]}

Trogolaphysa fuelleborni (Börner, 1903) n. comb. Paronella fuelleborni Börner, 1903: 143-144, fig. 10 (Langenburg, Nyassa region, Tanzania). — Wahlgren, 1908 (compared to P. nigromaculata, description of color pattern and claw structure, Kilimanjaro, Tanzania). — Philiptschenko 1926: 192-193 (proposed as synonym of P. nigromaculata Schött, 1903). — Denis 1933: 263, 268 (compared to T. separata Denis, 1933, suggestion that synonymization with P. nigromaculata by Philiptschenko [1926] is unjustified). — Womersley 1934: 441, 471 (records from literature?, Tanzania [Nyassa, Kilimanjaro], Madagascar?). — Martynova 1961: 482-483 (synonym of P. nigromaculata). Dicranocentruga fuelleborni – Mitra 2002c: 114. TYPE LOCALITY. — Langenburg (now Tukuyu), Nyassa region, Tanzania. DISTRIBUTION. — Tanzania (Tukuyu, Kilimanjaro), Madagascar? REMARKS This species was originally described from southern Tanzania, near the northern shore of Lake Malawi. Börner’s (1903) original description dealt mostly with generalized characters that provide little valuable information to characterize the species. This has resulted in general confusion on the identity of the species, and whether it is a junior synonym of P.nigromaculata Schött, 1903. Thus, on examining specimens from Kenya he identified as P. nigromaculata, Philiptschenko (1926) declared the species to be so variable as to encompass the morphology observed in T. fuelleborni n. comb. Denis (1933) included T. fuelleborni n. comb. in his key to Paronella species and pointed out that the placement of inner teeth of the unguis (on the distal half of the inner margin) distinguishes this species from P. nigromaculata. Denis further suggested that Philiptschenko’s material from Kenya may represent a different species altogether. Womersley (1934) published a list of African species which included a report of T. fuelleborni n. comb. from Madagascar. Womersley (1934) did not provide additional information about the material on which this report was based, and it remains unclear if the report from Madagascar is real or a mistake. Martynova (1961) supported Philiptschenko’s conclusion based on the variation displayed by material from Kenya she identified as P. nigromaculata. We place this species in Trogolaphysa based on its general similarity to P.nigromaculata (see Remarks to P. nigromaculata below). Given the confusion in the literature as to how to characterize T. fuelleborni n. comb., it is difficult to provide an adequate description without examining type specimens or topotypical material. Following Börner’s (1903) original description the species has a light brownish yellow background color with dark blue pigment covering flanks of body, head darker anteriorly, 1 st and 2 nd antennal segments light, all other antennal segments dark blue, coxae dark blue; tenet hair long and spatulate; unguis with 4 inner teeth, paired teeth subequal, inserted on distal half of inner edge, proximal unpaired tooth shorter than paired teeth and inserted on distal half of inner edge, away from basal teeth; dens with 1 row of spines; mucro short, with 4 teeth. Based on the descriptions available, the only character distinguishing T. fuelleborni n. comb. is the displacement of the inner teeth of the unguis to the distal half of the claw (Fig. 5C).

KEY TO AFRICAN SPECIES OF PARONELLA -LIKE FORMS HERE ASSIGNED TO TROGOLAPHYSA MILLS, 1938 1. Unguiculus truncate (Fig. 5H).................................................................................................................... 2 — Unguiculus lanceolate (Figs 5 B-E)............................................................................................................... 3 2. Metathorax to 2 nd abdominal segment uniformly dark blue (Fig. 5J); dens with inner row spines apically acuminate (Ivory Coast)............................................................ T. beguei (Delamare Debouteville, 1951) n. comb. — Mesothorax to 2 nd abdominal segment never uniformly dark blue; dens with inner row spines apically asymmetrically ciliate, brush-like (Cameroon, Ivory Coast, Gabon)............. T. penicillata (Schött, 1927) n. comb. 3. Mucro with 5 teeth (Fig. 5F)....................................................................................................................... 4 — Mucro with 3-4 teeth (Figs 3C, D, 5G)....................................................................................................... 5 4. Abdomen 4 medially with dark blue square; unguis with one paired inner tooth clearly larger than other, much larger than proximal unpaired tooth; Mucro with basal teeth more or less evenly distributed (Fig. 5F) (Gabon).............................................................................................................. T. monomaculata (Barra, 1969) — Abdomen 4 white, without marks; Unguis with paired inner teeth subequal, and not much larger than unpaired tooth; Mucro with basal teeth grouped on proximal end (Fig. 5G) (Gabon).............. T. vivieni (Barra, 1969) 5. Mucro with 3 teeth; tenet hair relatively short in relation to unguiculus (Fig. 5A); Unguis with 3 inner teeth (Ivory Coast).................................................................... T. grassei (Delamare Debouteville, 1952) n. comb. — Mucro with 4 teeth; tenet hair normally long in relation to unguiculus (Fig. 5B); Unguis with 3-4 inner teeth.... 6 6. Unguis with 4 inner teeth............................................................................................................................ 7 — Unguis with 3 inner teeth.......................................................................................................................... 10 7. Antennomere 1 white, antennomeres 2-4 uniformly dark blue (Fig. 5I); femora white, tibiotarsi blue (Guinea)........................................................................ T. lamottei (Delamare Debouteville, 1950) n. comb. — Antennomeres 1-2 white/yellow or light blue, antennomeres 3-4 darker; legs with all segments blue, brown or yellow.......................................................................................................................................................... 8 8. Unguis with all teeth inserted on distal half of inner edge (Fig. 5C) (Tanzania, Madagascar?)................................................................................................................................... T. fuelleborni (Börner, 1903) n. comb. — Unguis with basal teeth inserted on basal half & unpaired teeth on distal half of inner edge (Fig. 5B)......... 9 9. Body yellow (Republic of Congo).................................................... T. ghesquierei (Marlier, 1945) n. comb. — Body blue (Madagascar).................................................................................................. T. judithnajtae n. sp. 10. Head macrochaetae A3 and Pa5 absent (Cameroon, Ivory Coast, Rwanda, Burundi, Tanzania, Ethiopia)..................................................................................................... T. nigromaculata (Schött, 1903) n. comb. — Head macrochaetae A3 and Pa5 present (Fig. 2A)...................................................................................... 11 11. Antennae, legs and body uniformly white; eye patch with up to 9 chaetae; Setiform labral papillae present; Dens with 1 row of spines (Kenya)................................................................. T. wahlgreni (Mitra, 2002) n. comb. — Antennae and legs uniformly dark blue, body with a general blue tint; eye patch with up to 6 chaetae; Setiform labral papillae absent; Dens with 2 row of spines (Madagascar)...................................... T. judithnajtae n. sp.

Trogolaphysa judithnajtae n. sp. (Figs 1-3) urn:lsid:zoobank.org:act: 16B525F2-9F21-4A3C-A6C7-F85DCDF23BFF TYPE MATERIAL. — Holotype. ♀, slide-mounted, Madagascar, Province d’Antsiranana, Nosy Be Réserve naturelle intégrale de Lokobe. 6.3 km ESE Hellville, 19-24.III.2001, 13°25’10”S, 48°19’52”E, pitfall trap in rainforests, B. L. Fisher, Griswold et al., CAS, coll. code BLF3418, CASLOT 041427. Paratypes. All deposited at CAS, unless otherwise specified. 4 in preparation (2 MNHN, 2 FSCA) and 8 (2 MNHN, 2 FSCA) in alcohol with same collection information as holotype; 5 slide mounted (1 MNHN, 1 FSCA) and 8 individuals in alcohol, Toliara Prov. Reserve Price Berenty, forêt de Bealoka, Mandraré River, 14.6 km 329°NNW Amboasary, Elev. 35 m, 3-8.II.2002, 24°57’25”S, 46°16’17”E, B. L. Fisher, Griswold et al., pitfall trap in gallery forest, Coll. code BLF5314, CASLOT 041; 5 slide mounted and up to 100 in alcohol, Toliara Prov. Reserve Price Berenty, forêt de Bealoka, Mandrake River, 14.6 km 329°NNW Amboasary, Elev 35 m., 3-8.II.2002, 24°57’25”S, 46°16’17”E, B. L. Fisher, Griswold et al. pitfall trap in Gallery Forest, code BLF3314, CASLOT 044128; 4 slide mounted and 16 in alcohol (2 FSCA), Antsiranana, forêt de Binara, 7.5 km 230°SW Daraina, Elev. 375m, 1.XII.2003, 13°15’18”S, 049°37’00”E, B. L. Fisher, pitfall trap, tropical dry forest, code BLF9558, CASLOT 041408; 5 slide mounted (1 FSCA) and 10 in alcohol, Mahajanga, Prov, Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, Elev. 150 m, 16-20.XI.2001, 19°42’34”S, 44°43’5”E, B. L. Fisher, Griswold et al., pitfall trap tropical dry forest on Tsingy, Coll. code BLF4433, CASLOT 04196; 5 slide mounted (1 FSCA) and 210 in alcohol (10 FSCA), Mahajanga Prov, Parc National Tsingy de Bemaraha, 3.4 km 93°E Bekopaka, Tombeau Vazimba, Elev. 50m, 6-10.XI.2001, 19°8’31”S, 44°49’41”E, B. L. Fisher, Griswold et al., pitfall trap in tropical dry forest, Coll. code BLF4230, CASLOT 041849; 3 slide mounted and 3 in alcohol, Toamansina Parcelle E 3 Tampolo, Elev. 10 m, 14.IV.2004, 17°17’00”S, 049°26’00”E, coll.: Malagasy ant team, yellow pan trap, littoral forest, Coll. code BLF10730, CASLOT 038816; 2 slide mounted and 350 in alcohol (10 FSCA), Province de Mahajanga, Parc national d’Ankarafantsika, forêt de Tsimaloto, 18.3 km 46°NE de Tsaramandroso, Elev. 135m, 2-8. IV.2001, 16°13’41”S, 46°8’37”E, B. L. Fisher, Griswold et al., pitfall trap in tropical dry forest, Coll. code BLF3597, CASLOT 044347; 4 slide mounted (1 slide FSCA) and 2 in alcohol, Toliara Prov., forêt de Mite, 20.8 km 29°WNW Tongobory, Elev. 75m, 23°31’27”S, 44°7’17”E, 27.II.-3.III.2002, B. L. Fisher, Griswold et al., pitfall trap in gallery forest, Coll. code BLF5848, CASLOT 041967. DISTRIBUTION. — Madagascar (Fig. 4). ETYMOLOGY. — The epithet honors Judith Najt, for her contributions to our understanding of the taxonomy of Paronellini. DIAGNOSIS. — Trogolaphysa judithnajtae n. sp. is the only blue species with six paired anterior head macrochaetae in the combination A2, A3, A5, M2, S3 and S5 that also has four inner macrochaetae on 4th abdominal segment (A3, A5, B4, B5) and labrum lacking distal margin denticles. DESCRIPTION Size Adults 2.3-2.6 mm long. Color pattern Background color in alcohol preserved individuals beige with blue pigment on all antennal and leg segments, manubrium, and base of collophore; interantennal field on head as dark as antennae; head light brown or uniformly blue, body mostly blue, eye patch black, anterior margin of Abd. 4 with unpigmented areas forming longitudinal streaks (Fig. 1). Scale distribution Present dorsally and ventrally on 1 st and 2 nd antennal segments but on 3 rd antennal segment limited to basal ¼ of dorsal face; dorsally and ventrally on head; dorsally on body; ventrally on furcula; absent from legs and collophore. Head Antennae up to 2.7× head length. Subapical sense organ of 4 th antennal segment a short rod contained in circular depression, guard sensillum rela tively short. Sense organ of 3 rd antennal segment with 2 main sensilla diamond shaped and inserted in a shallow depression. Eyes 8+ 8, usually appearing as 6+6, with eyes H and G reduced as typical for genus, clearly visible in few individuals; eye valley usually with 6 (5-7) ciliate chaetae (Fig. 2B). Head with 6 (5.7) anterior paired macrochaetae (A2, A3, A5, M2, S3, S5), macrochaeta S4 sometimes present (arrow in Fig. 2A), single individual with 5 macrochaetae lacks M2 and S4; macrochaeta An3p1i present; paired posterior macrochaetae Pa5 and Pm3 present. Pre-labral chaetae ciliate, all labral chaetae smooth. Distal margin of labrum smooth, without denticles. Outer maxillary lobe with basal chaeta and apical appendage smooth and subequal; sublobular plate with 2 appendages. Labial triangle formula as M1M2rEL1L2A1-5 (Fig. 2C); r acuminate, smooth, shortest of posterior chaetae; all other posterior chaetae ciliate, L2 subequal to L1. Postlabial field with 4 +4 chaetae along cephalic groove, 1 chaeta in column C, and 5-6 chaetae between columns C and O. Body Body macrochaetae formula 6(5-7)0/0244+0+6. Mesothorax (Fig. 2D), with 1 anterior (a5) and 6 (5-7) posterior (p3 com- plex) macrochaetae. Metathorax with 4 inner microchaetae, without macrochaetae. Abdominal segment 1 with 3 inner microchaetae, chaeta a6 absent. Abdominal segment 4 (Fig. 2E) with pseudopore inserted in field posterior to T4; inner macrochaetae A3, A5, B4 and B5 present; A3 well anterior to T2; A5 inserted posterior to pseudopore; B4 inserted closer to A5 than to B5; 6 large lateral macrochaetae usually present, sometimes additional lateral macrochaetae present (Fig. 2F). Abdominal segment 4 with 10 +10 posterior chaetae; posterior intersegmental membrane with up to 6+ 6 lenticular organs as described by Jantarit et al. (2013) for Cyphoderopsis. Legs Trochanteral organ with up to 39 chaetae. Metathoracic claw complex as in Fig. 3A, B, tenent hair spatulate on all legs, sometimes weakly so, 1.64 × (n= 10) as long as unguiculus; claws usually with 3 (Fig. 3A), sometimes with 4 inner teeth (Fig. 3B), basal teeth subequal or one slightly longer; proximal unpaired tooth shorter than basal teeth; distal unpaired tooth inconspicuous; inner teeth spread for individuals with 3 teeth as 46%, 46%, 73%, for individuals with 4 teeth as 42%, 42%, 72%, 90%. Dorsal and lateral teeth ending on basal quarter of unguis, lateral teeth well developed. Unguiculus lanceolate, posterior edge smooth. Collophore Anterior face with 3 + 3 inner distal macrochaetae. Lateral chaetae up to 14 +14, with 4 distal chaetae smooth and sharply tapered. Posterior face distally with many ciliate chaetae. Furcula Dens with 2 rows of ciliated spine-like chaetae: inner row with up to 28 spines, outer row with up 18 spines. Mucro with 4 teeth, basal teeth subpaired (Fig. 3C), one individual with basal teeth grouped on one side (Fig. 3D); mucro subrectangular, 1.6-1.9 × as long as width of distal end of dens and 0.79× as long as inner margin of metathoracic unguis. REMARKS The new species is most similar to T. wahlgreni n. comb. from which it can be distinguished by color pattern, absence of labral denticles, and number of rows of spines on dens as detailed in Table 2. The morphology of the new species overlaps some of the extensive morphological variation reported for T. nigromaculata n. comb. (see below). Nevertheless, Mitra (2002c) points out that T. nigromaculata n. comb. lacks dorsal head macrochaetae A3 and Pa5, which are present in the new species. The few characters described for T. ghesquierei n. comb. make it almost indistinguishable from the new spe- cies, only the yellow body and brown legs in T. ghesquierei n. comb. separate the two forms. Womersley (1934) reported T. fuelleborni n. comb. from Madagascar, but none of the specimens we examined from across the island has the paired ungual teeth inserted on the distal half of the inner edge, as illustrated by Börner (1903). Based on Börner’s original drawing the spread of inner teeth in T. fuelleborni n. comb. is 56:56:80:90, whereas in the four-toothed form of T. judithnajtae n. sp. it is 42:42:72:90 The material examined includes populations from throughout the island, from Nosy Be and the forêt de Binara in the North to the forêt de Bealoka in Toliara Province in the Southeast (Fig. 4). At first, the variation on some important diagnostic character (number of dorsal head macrochaetae, number of macrochaetae in the p3 complex of the meso- thorax, claw dentition) lead us to think we were dealing with more than one species. We evaluated the distribution of characters among individuals and localities but did not find a correlation between them. The individuals with head macrochaeta S4 (i.e., seven paired anterior macrochaetae) or lacking macrochaeta A2 (five paired anterior macrochaetae) all come from the same population, bear three inner ungual teeth and do not differ from other individuals in the sample in any other character.Two southern populations (BLF4230 & BLF5848) are characterized by having four inner ungual teeth, but otherwise are identical to populations from the north in the number of macrochaetae and other diagnostic characters. Two individuals from the north, one each from localities BLF9558 & BLF3418 (type locality) also have four teethed-claws. Although we do not dismiss the possibility that some of these forms represent different species (i.e., T. judithnajtae n. sp. could be a species complex), currently we do not have evidence to circumscribe them unambiguously. Determination of the significance of morphological variation may have to await a molecular analysis., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on pages 184-188, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["JANTARIT S., SATASOOK C. & DEHARVENG L. 2013. - The genus Cyphoderopsis Carpenter (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae. Zootaxa 3721: 049 - 070. https: // doi. org / 10.11646 / zootaxa. 3721.1.2","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116.","WOMERSLEY H. 1934. - On some Collembola Arthropleona from South Africa and Southern Rhodesia. Annals of the South African Museum 3: 441 - 475.","BORNER C. 1903. - Uber neue Altweltliche Collembolen, nebst Bermerkungen zur Systematik der Isotominen und Entomobryinen. Gesellschaft naturforschender Freunde: 129 - 182."]}

Trogolaphysa lamottei (Delamare Debouteville, 1950) n. comb. Paronella lamottei Delamare Debouteville, 1950b: 43 -44, figs 1-6 (Mt Nimba, Guinea). Dicranocentruga lamottei – Mitra 2002c: 114. TYPE LOCALITY. — Mt Nimba, Guinea. DISTRIBUTION. — Guinea. REMARKS This species has not been reported again since the original description. The only additional relevant citation is that by Mitra (2002c), in which he transferred it to the genus Dicranocentruga. Paronella lamottei is placed in Trogolaphysa based on the shape of the mucro and absence of manubrial spines. However, this species approaches Paronella /Trychorypha atrofasciata in the presence of relatively long antennae and in having the manubrium distinctly longer than the dens, and it ultimately may belong to that species/genus group. Paronella lamottei is characterized by a long and thin body, with relatively long antennae, which is common in cave forms but atypical for surface Trogolaphysa; body white, except for lateral margins of the mesothorax to the 1 st abdominal segment and tibiotarsi; 1 st antennal segment white, but all other segments dark blue; tenet hair relatively long and strongly spatulate; unguis with 4 inner teeth; unguiculus lanceolate; dens with two rows of spines; and mucro with 4 teeth. This species should be easy to recognize by its general habitus and color pattern (Fig. 5I)., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on pages 183-184, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["DELAMARE DEBOUTTEVILLE C. 1950 b. - Deux nouvelles especes de collemboles du Mont Nimba (Guinee francaise) recoltees par M. Maxime Lamotte. Bulletin de la Societe zoologique de France 75: 43 - 45.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116."]}

Trogolaphysa nigromaculata (Schött, 1903) n. comb. Paronella nigromaculata Schött, 1903: IV, VII, pl. 1 fig. 8-11 (Bonge, Cameroon). — Wahlgren 1908: 2, 4, 6-8 (description, material used by Mitra 2002c to describe D. wahlgreni, Meru, Kenya). — Philiptschenko 1926: 192-193 (description, senior synonym of P. fuelleborni, Molo, British East Africa). — Schött 1927: 25, 27-30, pl. II figs 10-15 (redescription and description of variation, key to species, Debundscha, Cameroon). — Handschin 1929: 77: 16, 22-23, fig. 5 (descriptive notes, Ethiopia). — Denis 1933: 267, 268 (key to species, comments on variation reported in literature). — Delamare Debouteville & Paulian 1952: 77 (mention, Ivory Coast). — Salmon 1956: 32-35, figs 88-102 (description, Rwanda, Burundi). — Paclt 1959: 56-58 (Neophorella dubia as junior synonym of T. nigromaculata, Natal, South Africa); 1967: 142 (record, Tanzania). — Martynova 1961: 848, 854-857, fig. 5-7 (descriptive notes on variation, Ethiopia). — Mitra 2002c: 106, 109, 112 (syntypes deposited at the Swedish Museum of Natural History, Stockholm, descriptive notes on chaetotaxy, comparison with D. whalgreni). Neophorella dubia Womersley, 1934: 464-465, fig. 12 (description, synonymyzed by Paclt 1959; Table Mount, Cape Town, South Africa). Dicranocentruga nigromaculata – Mitra 1992: 212. — Mitra 2002c: 114, pl. I, fig. 1D (photo of slide-mounted syntype). TYPE LOCALITY. — Bonge, Cameroon. DISTRIBUTION. — Cameroon, Ivory Coast, Rwanda, Burundi, Tanzania, Ethiopia. REMARKS The description of Neophorella dubia Womersley, 1934 was based on a single individual collected in Cape Town, South Africa. Neither the original description of the species nor the accompanying figures recall or resemble a paronellid by any stretch of the imagination (habitus isotomid-like, with 3 rd abdominal segment longer than 4 th abdominal segment, dens twice as long as manubrium, scales absent, and mucro with three apical teeth). However, Paclt (1959) studied the holotype and indicated that the claw does resemble that of T. nigromaculata n. comb., and that dental spines are present. Paclt (1959) did not mention the structure of the mucro. Ireson & Greenslade (1990) re-examined the type specimen (and only known individual) of N. dubia and concluded that the species belongs in Tomoceridae Schäffer, 1896, as originally proposed by Womersley. Additional attempts to find the species at the type locality have been unsuccessful (Janion-Scheepers et al. 2015), but it seems safe to conclude that N. dubia does not belong in Trogolaphysa. Trogolaphysa nigromaculata n. comb. is transferred to Tro - golaphysa on account of Mitra (2002c), who examined some types (see below) and placed the species in Dicranocentruga, a junior synonym of Trogolahysa (Soto-Adames &Taylor 2013). Trogolaphysa nigromaculata n. comb. is the most often reported and cited member of the genus in Africa, with putative distribution covering most of the tropical and southern subtropical regions of the continent. The original and subsequent descriptions by Schött (1903, 1927), although relatively extensive, provide few characters useful in distinguishing the widespread populations. The absence of a description of relevant diagnostic characters of the type material has lead authors to consider T. nigromaculata n. comb. a very variable species. The main diagnostic character of the species is related to the morphology of the unguis, which Schött (1903) described as having three inner teeth. However, samples including individuals with three or four inner ungual teeth was interpreted by various authors as intraspecific variation. Following this interpretation in claw variation the species was reported from Kenya (Whalgren 1908), Ethiopia (Philiptschenko 1926; Handschin 1929; Martynova 1961), Ivory Coast (Delamare Debouteville & Paulian 1952), Rwanda and Burundi (Salmon 1956), and Tanzania (Paclt 1967). It is unlikely that populations from such distant localities (which cover more than 7000 km) and diverse environmental conditions (from the lowland tropical rain forest of Ivory Coast and Cameroon, to the highland forest of Rwanda and Burundi, and the xeric conditions of Ethiopia) represent the same species. Mitra (2002c) studied some syntypes of T. nigromaculata n. comb. from Cameroon and individuals with three-toothed unguis from Kenya (originally identified as P. nigromaculata by Wahlgren 1908) and found the two populations to be easily distinguishable using head chaetotaxy. Extrapolating from Mitra’s (2002c) observation, it is likely that variation in claw morphology reported in the literature reflects a mixture of species and that T. nigromaculata n. comb. as currently delimited is a species complex. The abbreviated description that follows is based on the original description by Schött (1903) with a few details of head chaetotaxy reported by Mitra (2002c). The redescription by Schött (1927) is unreliable because the material on which it is based was collected in Debundscha, not in the type locality of Bonge. Schött (1927) reports two color forms, the pale one matching the original description, and a dark one with more extensive blue pigment distributed along the flanks. In addition to differences in color pattern, the unguiculus in the dark form is truncate instead of lanceolate, and one of the inner paired ungual tooth is considerably longer than the other. The dark form is likely an unnamed species (Mitra 2002c might have been referring to these two forms when he pointed out that Schött’s material from Cameroon and Wahlgren’s material from Kenya includes three species). The light form of T. nigromaculata n. comb. is characterized by a whitish body, with dark blue pigment on all antennal segments, eye patch and coxae; head dorsally with five paired anterior macrochaetae (A2, A5, M2, S3, S5), paired posterior macrochaeta Pa5 absent; tenet hair somewhat short and spatulate; unguis with three inner teeth, basal teeth inserted on basal half of inner edge, unpaired tooth on distal half; unguiculus lanceolate; mucro square, quadridentate, with basal teeth paired., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on pages 189-190, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["SCHOTT H. 1903. - Uber zwei Gattungen der Apterygoten Insekten. Redogorelse for Allamanna Larvorken I Linkoping. Esko och Vadstena. Underlarsaret: 3 - 16.","WAHLGREN E. 1908. - Wissensch Ergebn. der Schwed Zoolog. Exp nach dem Kilimandjaro, dem Meru 1905 - 1906. Unter Leitung von Prof. Dr. Yngve Sjostedt. 18: Apterygogenea 1. Collembola: 1 - 10.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116.","PHILIPTSCHENKO J. 1926. - On the Collembola collected by the expedition of V. A. Dogiel and I. I. Sokolov in British East Africa. Revue russe d'Entomologie 20: 180 - 196.","SCHOTT H. 1927. - Kamerunische Collembola. Medd. Linkopings Hogre Allmanna Laroverk redogorelse: 1 - 39.","HANDSCHIN E. 1929. - Collembola from Abyssinia. Transactions of the Entomological Society of London 77: 15 - 28. https: // doi. org / 10.1111 / j. 1365 - 2311.1929. tb 00678. x","DENIS J. R. 1933. - Contributo alla conoscenza del \" microgenton \" di Costa Rica, III. Collemboles de Costa Rica avec une contribution au species de l'ordre (deuxieme note). Bolletino del Laboratorio di Zoologia Generale e Agraria Portici 27: 222 - 322.","SALMON J. T. 1956. - Contribution a l'etude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky 1953): LXXIX. Collembola. Sciences zoologiques 51: 9 - 40.","PACLT J. 1959. - Collembola. South African Animal Life 6: 24 - 78.","MARTYNOVA E. F. 1961. - Materials on the fauna of Collembola of Ethiopia. Revue d'Entomologie de l'USSR 40: 848 - 857.","WOMERSLEY H. 1934. - On some Collembola Arthropleona from South Africa and Southern Rhodesia. Annals of the South African Museum 3: 441 - 475.","MITRA S. K. 1992. - Fixation of the concept of Paronella Schott, 1893 [Collembola: Entomobryidae]. Records of the Zoological Survey of India 92: 211 - 224.","IRESON J. E. & GREENSLADE P. 1990. - Lasiofinus gen. n. (Collembola: Tomoceridae) from Tasmania and re-examination of Neophorella dubia Womersley (Tomoceridae). Journal of the Australian Entomological Society 29: 205 - 214. https: // doi. org / 10.1111 / j. 1440 - 6055.1990. tb 00350. x","PACLT J. 1967. - On South and Central African Collembola. Journal of the Entomological Society of South Africa 29: 135 - 147."]}

Trogolaphysa penicillata (Schött, 1927) n. comb. Paronella penicillata Schött, 1927: 29-30, pl. II, fig. 16-20, key (Debundscha, Cameroon). — Denis 1933: 268, key. — Delamare Debouteville 1947: 453; 1948: 316, 421 (association with termite Noditermes curvatus, Ivory Coast); 1951b: 205, 209, 212, fig. 51 (Ivory Coast). — Delamare Debouteville in Delamare Debouteville & Paulian 1952: 77 (Le Banco, Ivory Coast). — Barra 1969: 189 (record, Ipassa, Gabon). — Mitra 2002c: 106 (syntypes deposited at the Swedish Museum of Natural History, Stockholm). Dicranocentruga penicillata – Mitra 1992: 212; 2002c: 114, 106, 114, fig. 3A (generic placement, incorrectly stating new combination, photo of syntype). TYPE LOCALITY. — Debundscha, Cameroon. DISTRIBUTION. — Cameroon, Ivory Coast, Gabon. REMARKS This species is placed in Trogolaphysa based on the absence of manubrial spines and the shape of the mucro. This species is restricted to Western Africa and is characterized by having a white background color, with blue pigment on lateral margin of the metathorax and 1 st abdominal segment, and a transversal band on anterior half of the 4 th abdominal segment; antennomere 1 light blue, all other antennal segments dark blue; tenet hair relatively long, and spatulate; unguis with 4 inner teeth, paired teeth inserted on basal half of inner edge; unpaired teeth shorter that paired teeth, inserted on distal half, far away from basal teeth; unguiculus truncate; dens with distal spines on inner row apically asymmetric and paintbrush-like; mucro with 4 teeth, basal teeth paired. This species is easy to recognize by its distinctive color pattern, truncate unguiculus and the paintbrush-like distal dental spines., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on pages 190-192, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["SCHOTT H. 1927. - Kamerunische Collembola. Medd. Linkopings Hogre Allmanna Laroverk redogorelse: 1 - 39.","DENIS J. R. 1933. - Contributo alla conoscenza del \" microgenton \" di Costa Rica, III. Collemboles de Costa Rica avec une contribution au species de l'ordre (deuxieme note). Bolletino del Laboratorio di Zoologia Generale e Agraria Portici 27: 222 - 322.","DELAMARE DEBOUTTEVILLE C. 1947. - Cycle evolutif d'un peuplement nidicole en Basse Cote-d'Ivoire. Bulletin du Museum 19: 453 - 455.","DELAMARE DEBOUTTEVILLE C. 1948. - Recherches sur les Collemboles termitophiles et myrmecophiles (Ecologie, Ethologie, Systematique). Archives de Zoologie experimentale et generale 85: 265 - 425.","DELAMARE DEBOUTTEVILLE C. 1951 b. - Microfaune du sol des pays temperes et tropicaux. Actualites scientifiques et industrielles 1160: 1 - 360.","DELAMARE DEBOUTTEVILLE C. & PAULIAN R. 1952. - Recherches sur la faune des nids et des terriers en Basse Cote d'Ivoire. Chapitre VIII, Etude systematique. I. - Collembola. Encyclopedie biogeographique et ecologique 8: 62 - 77","BARRA J. - A. 1969. - Collemboles du Gabon. Revue Biologia Gabonica 5: 189 - 216.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116.","MITRA S. K. 1992. - Fixation of the concept of Paronella Schott, 1893 [Collembola: Entomobryidae]. Records of the Zoological Survey of India 92: 211 - 224.","MARLIER G. 1945. - Collemboles du Congo belge. Revue de Zoologie et de Botanique africaines 38: 252 - 264.","BORNER C. 1903. - Uber neue Altweltliche Collembolen, nebst Bermerkungen zur Systematik der Isotominen und Entomobryinen. Gesellschaft naturforschender Freunde: 129 - 182.","CARPENTER G. H. 1916. - The Apterygota of the Seychelles. Proceedings of the Royal Irish Academy 33: 1 - 108.","DELAMARE DEBOUTTEVILLE C. 1951 a. - Nouveaux Paronelliens de la Cote d'Ivoire. Bulletin de l'Institut francaise d'Afrique noire 13: 1072 - 1075.","DELAMARE DEBOUTTEVILLE C. 1950 b. - Deux nouvelles especes de collemboles du Mont Nimba (Guinee francaise) recoltees par M. Maxime Lamotte. Bulletin de la Societe zoologique de France 75: 43 - 45."]}

Trogolaphysa vivieni (Barra, 1969) Paronella (Microparonella) vivieni Barra, 1969: 189, 207-212, fig. 6A-N (Ipassa, Gabon). Lepidonella vivieni – Deharveng & Bedos 1995: 22. Trogolaphysa vivieni – Soto-Adames & Bellini 2015: 337. REMARKS Like T. monomaculata, T. vivieni was originally described in Microparonella, transferred to Lepidonella by Deharveng & Bedos (1995) and moved to Trogolaphysa by Soto-Adames & Bellini (2015). Trogolaphysa vivieni is characterized by a uniformly whitish body, with characteristic blue pigment on the basal half of antennomeres 2-3, all antennomere 4 and metathoracic trochanter; scales present on antennomeres 1-3 and on coxae, trochanter, and femur of all legs; tenet hair long and weakly spatulate; unguis with 3 inner teeth, paired teeth subequal; unguiculus lanceolate; dens with two rows of spines; mucro long, with 5 teeth, apical tooth much longer than all basal teeth. This is the only white African species with dark blue pigment limited to the antennae and metatrochanter, and a five toothed mucro with a distinctly long apical tooth., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 192, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["BARRA J. - A. 1969. - Collemboles du Gabon. Revue Biologia Gabonica 5: 189 - 216.","DEHARVENG L. & BEDOS A. 1995. - Lepidonella lecongkieti n. sp., premiere Collembole cavernicole de Vietnam. Bulletin de la Societe entomologique de France 100: 21 - 24.","SOTO- ADAMES F. N. & BELLINI B. C. 2015. - Dorsal Chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea). Florida Entomologist 98: 330 - 341. https: // doi. org / 10.1653 / 024.098.0152"]}

Trogolaphysa wahlgreni (Mitra, 2002) n. comb. Dicranocentruga wahlgreni Mitra, 2002c: 109-113, figs 1-3, 5 (Meru, Kenya). Paronella nigromaculata – Wahlgren 1908: 2, 4, 6-8 (wrong identification). REMARKS Mitra (2002c) described this species based on material Wahlgren (1908) identified as P. nigromaculata. This species is characterized by being uniformly yellow or white, with pigment only on eye patch; labral papillae present; anterior head chaetotaxy with one unpaired (A0) and six paired (A2, A3, A5, M2, S3, S5) macrochaetae, posterior chaetotaxy with one paired macrochaeta (Pa5); mesothorax to 4 th abdominal segment with 60/0114 inner macrochaetae; tenet hair long, strongly spatulate; unguis with 3 inner teeth, paired teeth inserted near middle of claw, one paired tooth slightly longer than other; unguiculus lanceolate; dens with one row of spines; mucro square, quadridentate. This is a typical Trogolaphysa, and very similar to T. judithnajtae n. sp. (see diagnosis for that species). The original body macrochaetae map for abdominal segments 2-4 published by Mitra (2002c: 113) confounds symbols for macrochaetae and pseudopores, but once the confusion is resolved, the inner chaetotaxy is typical Trogolaphysa. This is the only African species uniformly white or yellow having head macrochaetae M2 and S5, labral papillae, unguis with three inner teeth and one row of spines on dens., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 192, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["MITRA S. K. 2002 a. - Status of Microparonella Carpenter, 1916 (Collembola: Entomobriyidae: Paronellinae). Records of Zoological Survey of India 100: 67 - 72.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116.","WAHLGREN E. 1908. - Wissensch Ergebn. der Schwed Zoolog. Exp nach dem Kilimandjaro, dem Meru 1905 - 1906. Unter Leitung von Prof. Dr. Yngve Sjostedt. 18: Apterygogenea 1. Collembola: 1 - 10."]}

Trogolaphysa ghesquierei (Marlier, 1945) n. comb. Paronella ghesquierei Marlier, 1945: 260, 261, fig. 6 (Eala, Belgian Congo). Dicranocentruga ghesourieri – Mitra 2002c: 114; lapsus calami. TYPE LOCALITY. — Eala, Democratic Republic of Congo. DISTRIBUTION. — Democratic Republic of Congo. REMARKS The only citations for this form are the original description, and its transfer to Dicranocentruga by Mitra (2002c). The species remains poorly characterized, but the shape of the mucro and absence of manubrial spines suggest it should be placed in Trogolaphysa. The Congolese species was described as uniformly yellow, with brown legs; a long spatulate tenent hair; unguis with four inner teeth; lanceolate unguiculus; dens with one row of spines; and four-toothed mucro. Among African Trogolaphysa, this appears to be the only uniformly yellow species with four inner ungual teeth and paired teeth inserted on the basal half of the claw., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 183, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["MARLIER G. 1945. - Collemboles du Congo belge. Revue de Zoologie et de Botanique africaines 38: 252 - 264.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116."]}

Trogolaphysa beguei (Delamare Debouteville, 1951) n. comb. Paronella beguei Delamare Debouteville 1951a: 1074, figs 5-12 (Ivory Coast); 1951b: 60, 82, 135, 166, 176, 254, 266, fig. 21 (ecology, Ivory Coast). Paronella beguei – Delamare Debouteville 1948: 309, 376, 421, nomen nudum. Dicranocentruga beguei – Mitra 2002c: 114. TYPE LOCALITY. — Le Banco, Ivory Coast. DISTRIBUTION. — Ivory Coast. REMARKS Delamare Debouteville first used the combination Paronella beguei in 1948 referring to a paronellid found together with termites. This first mention of P. beguei included neither a morphological description nor drawings and therefore it is a nomen nudum. The only other significant citations for P. beguei are those of Delamare Debouteville (1951b), concerning its ecology, and Mitra (2002c) in which the author transfers the species to the genus Dicranocentruga Wray, 1953. We assign this species to Trogolaphysa based on mucro shape, which is typical for the genus, short and bearing 4 teeth, and the relatively long antennae. Otherwise, this is a well-marked species, unique among African forms in having a distinctive color pattern (Fig. 5J) and truncate unguiculus (Fig. 5H)., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 182, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["DELAMARE DEBOUTTEVILLE C. 1951 a. - Nouveaux Paronelliens de la Cote d'Ivoire. Bulletin de l'Institut francaise d'Afrique noire 13: 1072 - 1075.","DELAMARE DEBOUTTEVILLE C. 1948. - Recherches sur les Collemboles termitophiles et myrmecophiles (Ecologie, Ethologie, Systematique). Archives de Zoologie experimentale et generale 85: 265 - 425.","MITRA S. K. 2002 c. - Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105 - 116."]}

Trogolaphysa monomaculata (Barra, 1969) Paronella (Microparonella) monomaculata Barra, 1969: 189, 212- 215, fig. 7A-K (Ipassa, Gabon). Lepidonella monomaculata – Deharveng & Bedos 1995: 22. Trogolaphysa monomaculata – Soto-Adames & Bellini 2015: 337. TYPE LOCALITY. — Ipassa, Gabon. DISTRIBUTION. — Gabon. REMARKS Barra (1969) described this species in the subgenus Microparonella. Deharveng & Bedos (1995) transferred all species assigned to Microparonella to the genus Lepidonella, but did not comment on the status of L. monomaculata. Soto-Adames & Bellini (2015) transferred the species to Trogolaphysa based on the chaetotaxy (2, 3, 3 bothriotricha on abdominal segments 2-4 and absence of head chaeta Ps 3) illustrated by Barra (1969). The habitus (Barra 1969: fig. 7A), mucro (fig. 7J, K) and claw shape (fig. 7G, H, I) suggest the species was described based on immature specimens. Trogolaphysa monomaculata is characterized by a uniformly light blue body, antennae, head, and legs, with a distinctive dark blue square band on 4 th abdominal segment; scales present on antennal segments 1-3 and on coxa, trochanter and femur of all legs; tenet hair long and weakly spatulate; unguis with 3 inner teeth, one paired tooth clearly larger than the other (Fig. 5D); unguiculus lanceolate; dens with two rows of ciliated spines; mucro rectangular, long, with 5 teeth. This species is unique among African forms in having a broad band on 4 th abdominal segment and a long, fivetoothed mucro with teeth evenly distributed throughout its length., Published as part of Nguyen, Minh & Soto-Adames, Felipe N., 2018, Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar, pp. 179-196 in Zoosystema 40 (10) on page 188, DOI: 10.5252/zoosystema2018v40a10, http://zenodo.org/record/3738229, {"references":["BARRA J. - A. 1969. - Collemboles du Gabon. Revue Biologia Gabonica 5: 189 - 216.","DEHARVENG L. & BEDOS A. 1995. - Lepidonella lecongkieti n. sp., premiere Collembole cavernicole de Vietnam. Bulletin de la Societe entomologique de France 100: 21 - 24.","SOTO- ADAMES F. N. & BELLINI B. C. 2015. - Dorsal Chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea). Florida Entomologist 98: 330 - 341. https: // doi. org / 10.1653 / 024.098.0152"]}

Nguyen, Minh, Soto-Adames, Felipe N. (2018): Annotated checklist of Afrotropical Trogolaphysa Mills, 1938 (Hexapoda: Collembola: Paronellidae) and description of a new species from Madagascar. Zoosystema 40 (10): 179-196, DOI: 10.5252/zoosystema2018v40a10

Paleonura borincana sp. nov. Figs. 1���6, Tables. 1���2 Type material. Holotype: female, microscopic slide preparation (INHS Insect Collection acc. # 551,630), Puerto Rico: Maricao, Monte del Estado Commonwealth Forest, Road PR-120, N 18.14396, W 66.97789, leaf litter in woods across entrance to old recreational area; 22.vi.1999. F. Soto-Adames, coll. Paratypes: one female (INHS acc. #551, 631), on microscope slide with same collection information as holotype; one male and one female (INHS acc. #���s 551, 632 and 551,633, respectively), microscope slide mounted, Puerto Rico: Caribbean National Forest, Pico del Oeste, 820���880 m above sea level, N 18.28, W 65.77, sierra palm forest litter associated with dwarf forest, 24.i.2001, B.A. Richardson & M.J. Richardson, coll. The holotype and one paratype are deposited in the insect collection of the Illinois Natural History Survey (INHS). Two paratypes remain in the collection of the senior author at the Universidad Nacional Aut��noma de M��xico, Mexico City. Type locality. Puerto Rico: Maricao, Commonwealth Forest. Etymology. Demonym for Borinquen, Spanish language corruption of the original Native American name for the island of Puerto Rico. Description. Length, 1.6 mm (n = 4). Color in alcohol white. Granulation fine, smaller than eye diameter. Only dorso-external tubercles on Abd. IV and V well developed. Body setae comprising microsetae; thick, hyaline, capitate macrosetae, and acuminate macrosetae, in addition to sensorial setae (Fig. 2). Ant. I with 7 setae, 3 dorsal with blunt tips. Ant. II with 11 setae, dorsal setae thicker and longer than ventral setae, 2 with blunt tips. Ant. III with 1 ventral microsensillum; sensorial organ with 2 globular sensilla in a cuticular fold and 2 guard sensilla; S.g. v. almost straight and longer than S.g.d. Ant. IV typical for the genus (Fig. 1), with all sensilla subequal in size and shape, 12 long acuminate setae, 1 small setae (i), 1 organite, and without subapical bulb. Ventrally on Ant. IV with setae br 3,4,5 and 6, and 3 cp (not drawn). Eyes 2 + 2. Mandibles with two thin and long teeth, maxillae styliform. Head without well developed tubercles, chaetotaxy as in Fig. 2. Labium with 9 pairs of setae. Ventral chaetotaxy of head with four setae Vea, 2 setae Vem and 6 setae Vep (not illustrated). Chaetotaxy of legs 1���3 as follows: coxae with 3,7,7 setae; all trochanters with 5 setae; femora with 12,12, 11 setae; tibiotarsi with 19,19, and 18 setae (Fig. 3, seta M not shown). Unguis without teeth. Thoracic and abdominal dorsal chaetotaxy in Figs. 2 and 4, dorsal tubercles not well developed. Group of setae De on Abd. V only with one sensorial seta (Fig. 5). Chaetotaxy by demitergite as in Table 1. Ventral tube with 4 + 4 setae. Furcal vestige with only 3 setae. Ventral chaetotaxy reduced as shown in Figure 6 and on Table 1. Furcula vestige with 3 setae. Female genital plate with 3+3 pregenital, 8���9 circumgenital and 2 eugenital setae. Each lateral anal tubercle with 12 setae and 2 microsetae, posterior one with one pair of setae. Abdominal sternites with strong chaetotaxy reduction (Fig. 6 and Table 1). Variation. One individual from Maricao has one additional supernumerary seta on head tubercle De. Remarks. Paleonura borincana sp. nov. is unique among the members of the genus in having the combination of cephalic setae A and B, two ocular setae and the absence of setae on the dorso-extenal tubercle of Abd. IV (only the sensorial seta remains). It resembles P. carayoni Massoud & Thibaud, 1987, from Guadalupe Island, but this species has only one ocular seta and 4 setae on the Di-De fused tubercle of Abd. V, while the new species has only 3 setae, in addition, Paleonura borincana sp. nov. measures 1.6 mm and P. carayoni is, at most, 1.0 mm long. Postcephalic, ventral and leg chaetotaxy of Paleonura boricana sp. nov. Thorax Legs, Published as part of Soto-Adames, Felipe N., 2017, On the genus Paleonura (Collembola: Neanuridae: Neanurinae) from the Americas and description of a new species from Puerto Rico, pp. 388-394 in Zootaxa 4318 (2) on pages 389-392, DOI: 10.11646/zootaxa.4318.2.11, http://zenodo.org/record/886727, {"references":["Massoud, Z. & Thibaud, J. M. (1987) Les Collemboles des Petites Antilles IV-Neanuridae (suite et fin). Revue d'Ecologie et de Biologie du Sol, 24, 91 ¯ 98."]}

Key to the species of Paleonura from the Americas and two Eastern Pacific P. spectabilis and P. formosana 1. De area of Abdominal segment IV without any ordinary seta, only the sensorial seta................................ 2 - De area of abdominal segment IV with one to four setae plus the sensorial seta..................................... 3 2. Antennofrontal area with setae ABCD............................................ P. boricana sp. nov. Puerto Rico - Antennofrontal area with only setae B and D......... P. carayoni Massoud & Thibaud, 1987 (Guadeloupe, Lesser Antilles) 3. No eyes (or one per side) only one ocular seta......... P. miniseta Massoud & Thibaud, 1987 (Guadeloupe, Lesser Antilles) - Two eyes per side, with 2 or 3 ocular setae.................................................................. 4 4. Antennofrontal area with only setae B....................... P. insularum (Carpenter, 1935) (Marquesas Is., Polynesia) - Antennofrontal area with two or more pairs of setae.......................................................... 5 5. Two ocular setae...................................................................................... 6 - Three ocular setae..................................................................................... 8 6. De tubercle on Thorax II and III with 3 setae and one sensorial setae, cephalic antennofrontal area with seta O..................................................................................... P. formosana (Yosii, 1965) (Taiwan) - De tubercle on Thorax II and III with 2 setae and one sensorial setae, cephalic antennofrontal area without seta O........ 7 7. De tubercle on Abd. IV with two setae plus one sensorial seta, 4 pais of setae on the antennofrontal area................................................................................... P. bilinskii Paśnik & Weiner, 2013 (Ecuador) - De tubercle on Abd. IV with three setae plus one sensorial seta, 2 pais of setae on the antennofrontal area.................................................................................. P. nuda Cassagnau & Oliveira, 1990 (Brasil) 8. Cephalic antennofrontal area with four or five pairs of setae and seta O........................................... 9 - Cephalic antennofrontal area with three pairs of setae, without seta O........................................... 10 9. Cephalic antennofrontal area with four pairs of setae, De on Abd. V with 4 setae...................................................................................... P. petebellingeri Palacios-Vargas & Sim��n-Benito, 2007 (USA) - Cephalic antennofrontal area with five pairs of setae, De on Abd. V with 6 setae....................................................................................................... P. saproxylica Smolis et Kadej, 2014 (USA) 10. De tubercle on Abd. IV with 4 setae plus the sensorial seta....................... P. spectabilis Cassagnau, 1982 (Nepal) - De tubercle on Abd. IV with 3 1 seta plus the sensorial seta................................................... 11 11. Cephalic De tubercle with 3 setae.................................. P. louisi Thibaud & Najt, 1993 (French Polynesia) - Cephalic De tubercle with 3 setae....................................... P. ili Christiansen & Bellinger, 1992 Hawaii, Published as part of Soto-Adames, Felipe N., 2017, On the genus Paleonura (Collembola: Neanuridae: Neanurinae) from the Americas and description of a new species from Puerto Rico, pp. 388-394 in Zootaxa 4318 (2) on pages 392-393, DOI: 10.11646/zootaxa.4318.2.11, http://zenodo.org/record/886727, {"references":["Massoud, Z. & Thibaud, J. M. (1987) Les Collemboles des Petites Antilles IV-Neanuridae (suite et fin). Revue d'Ecologie et de Biologie du Sol, 24, 91 ¯ 98.","Carpenter, G. H. (1935) Marquesan Collembola. B. P. Bishop Museum Bulletin, 114, 363 - 78.","Yosii, R. (1965) On some Collembola of Japan and adjencent countries. Contributions from the Biological Laboratory Kyoto University, 19, 1 - 71.","Pasnik, G. & Weiner, W. M. (2013) Paleonura bilinskii (Collembola, Neanuridae, Paleonurini), a new species from Ecuador. Zootaxa, 3702 (3), 295 - 300. https: // doi. org / 10.11646 / zootaxa. 3702.3.7","Cassagnau, P. & Oliveira de, E. P. (1990) Les collemboles Neanurinae d'Amerique du Sud. Bulletin de la Societe D'histoire Naturelle de Toulouse, 126, 19 - 23.","Palacios-Vargas, J. G. & Simon Benito, J. C. (2007) A new genus and three new species of Neanuridae (Collembola) from North America. Journal of Cave and Karst Studies, 69, 318 ¯ 325.","Smolis, A. & Kadej, M. (2014) A new saproxylic Paleonurini (Collembola, Nenauridae) species from North America with the first record of Galanura agnieskae Smolis, 2000 from the Continent. Florida Entomologist, 97 (4), 1386 - 1394. https: // doi. org / 10.1653 / 024.097.0413","Cassagnau, P. (1982) Sur les Neanurinae primitives suceurs et les lignees qu'en derivent (Collemboles). Travaux du Laboratoire d'Ecobiologie des Arthropodes Edaphiques, Toulouse, 3, 1 - 11.","Thibaud, J. - M. & Najt, J. (1993) Les Collemboles (Insecta) de l'atoll de Fangataufa. Bulletin du Museum national d'Histoire naturelle, 4 e Serie, 15, 95 - 105.","Christiansen & Bellinger (1992) Insects of Hawaii. Volume 15. Collembola, Honolulu, 1 - 445."]}

Soto-Adames, Felipe N. (2017): On the genus Paleonura (Collembola: Neanuridae: Neanurinae) from the Americas and description of a new species from Puerto Rico. Zootaxa 4318 (2): 388-394, DOI: https://doi.org/10.11646/zootaxa.4318.2.11

Campylothorax dominicanus sp. nov. Figures 10 ��� 11 Etymology The epithet refers to the apparent widespread distribution of the species in the northeastern section of the Dominican Republic. Material examined Holotype, slide-mounted, Dominican Republic, La Vega, Cordillera Central, Loma Casabito, 15.8 km NW, Bonao, 19��2'11.976''N, 70��31'8.0034''W, 1455 m elevation, evergreen cloud forest, yellow pan trap, 28 May 2003, sample DR21262, J. Rawlins, C. Young, R. Davidson, C. N����ez, P. Acevedo; one paratype in preparation and one juvenile in alcohol same collection data as holotype; two indviduals in alcohol, Duarte, Reserva Loma Quita Espuela, Canelo, 13.2 km NNE San Francisco de Macoris, 19��24'46.044''N, 70��9'52.02''W, edge of broadleaf forest, yellow pan trap, 6 April 2006, sample DR11263, C. Young, R. Davidson, J. Rawlins; two individuals in alcohol, poor condition, Cordillera Central, Reserva Valle Nuevo, La Nevera, 15.1 km SE Valle Nuevo, 18��41'47.04''N, 70��35'30.0114''W, 2252 m elevation, montane meadow in cloud pine forest, yellow pan trap, 3 June 2003, sample DR24462, R. Davidson, C. Young, C. N����ez, J. Rawlins, P. Acevedo, M. de la Cruz. Size. Up to 2.2 mm. Colour pattern. Background colour white (Figure 10 A); dark purple pigment evenly distributed on basal two-thirds of antennae, head, body, legs and manubrium; most intense on posterior three-quarters of Abd. 4; anterior margin of Abd. 4 with short, irregular, white or light blue longitudinal stripes (Figure 10 B). Head. Dorsal head chaetotaxy as in C. sabanus (Figure 1 B) with eight Mc along antennal margin and four other Mc along inner eye patch margin; with one unpaired (A0) and seven paired anterior (A2, A3, A5, S2, S3, S4, S5), and one posterior (Pa5) Mc. Pre-labral chaetae smooth. Maxilla lamella 1 and 2 as in C. notidanus sp. nov. Lateral appendage of labial papilla E long, reaching tip of papilla. Number of chaeta along cephalic groove unclear, apparently six. Body. Body Mc as 74/0241+7+0+5. Th. 2 dorsally flat; Th. 3 rounded, relatively short, partially covering central section of Abd. 1. Th. 2 with seven Mc in p3 complex; Th. 3 with four Mc in p2 ��� 3 complex. Abd. 3 with 3 + 2 lateral mesochaetae as in C. notidanus sp. nov. Inner section of Abd. 4 with anterior triangle formed by paired A1 and medial unpaired Mc; inner posterior Mc 6 + 6 organised as in Figure 11 A; Mc A3 inserted posterior to pseudopore. Lateral section of Abd. 4 with five large Mc (E2, E3, E4, F1, F2) and three other, smaller latero-posterior Mc; E4 inserted anterior to D2; F3 absent; posterior chaetae 16 + 16. Legs. Trochanteral organ with up to 26 chaetae. All tenent hairs spatulate, relatively short, reaching down to level of paired inner ungual teeth (Figure 11 B). All claws with two inner teeth and two outer teeth: inner teeth subequal, inserted on basal half of all legs; lateral teeth short, often obscured, inserted on basal quarter of outer edge; dorsal tooth absent. Unguiculus of all legs truncate, with one inner tooth; unguiculus on fore and middle legs shorter than on hind legs. Furcula. Mucro typical for genus, 2.3 ��� 2.4 times as long as inner edge of hind claw. Remarks Campylothorax dominicanus sp. nov. is the only member of the genus with the combination of an evenly pigmented purple body and Abd. 4 with short anterior white stripes, Abd. 4 lateral Mc F3 absent, unguis with two inner teeth and tenent hair reaching only to the level of inner ungual teeth. Dark specimens of C. sabanus, C. cubanus Gruia, 1983 and C. mitrai are similar to C. dominicanus sp. nov., but the absence of Mc F2 and presence of only two inner ungual teeth distinguishes the new species. The pattern of white stripes on Abd. 4 is similar to that of C. longicornis Sch��tt, 1893 but the Cameroonian species has antero-medial multiplets (brow) on Abd. 4. This seems to be the most common species on the northeastern section of the Dominican Republic, and the only one found at low elevations., Published as part of Felipe N. Soto-Adames, 2016, Chaetotaxy of first-instar Campylothorax sabanus (Wray), and description of three new Campylothorax species from Hispaniola (Collembola, Paronellidae), pp. 1583-1612 in Journal of Natural History 50 (25) on pages 1602-1605, DOI: 10.1080/00222933.2016.1145272, http://zenodo.org/record/269880, {"references":["Gruia MM. 1983. Collemboles Arthropleones de Cuba recoltes par les expeditions cubano-roumaines en 1969 - 1973, II. Resultats des expeditions biospeleologiques cubano-roumaines a Cuba. 4: 191 - 205.","Schott H. 1893. Beitrage zur Kenntnis der Insektenfauna von Kameroon. I. Collembola. Bihang till Kongliga Svenska Vetenskaps-Akademien Handligar. 19: 1 - 28 + 7 plates."]}

Campylothorax notidanus sp. nov. Figures 4 ��� 6 Etymology The epithet refers to the greatly enlarged, conic metanotum. Material examined Holotype, on slide, Dominican Republic, La Vega, Cordillera Central, Loma Casabito, 15.8 km NW, Bonao, 19��2'11.976''N, 70��31'8.0034''W, 1455 m elevation, evergreen cloud forest, yellow pan trap, 28 May 2003, sample DR21262, J. Rawlins, C. Young, R. Davidson, C. N����ez, P. Acevedo; four paratypes on slides and 32 adults, subadults and juveniles in alcohol, same collection information as holotype; one paratype in alcohol, La Vega, Cordillera Central, 4.1 km SW El Convento, 19��2'15.5394''N, 70��42'48.024''W, 1730 m elevation, dense secondary evergreen forest, yellow pan trap, 31 May 2003, sample DR22262, J. Rawlins, R. Davidson, C. Young, C. N����ez and P. Acevedo. Size. Up to 4.0 mm. Colour pattern. Antennae white or lightly pigmented with purple (Figure 4 A); head light purple, Th. 2, anterio-lateral margins of Th. 3, coxae and trochanters dark purple; legs white distal to trochanter; Abd. 3 with separated medial and lateral spots or bands; Abd. 4 anteriorly with 1 ��� 3 paired narrow longitudinal stripes (Figure 4 A, C), sometimes reaching latero-medial patches; Abd. 4 triangular latero-medial patches (Figure 4 B, C) sometimes extending medio-posteriorly to join narrow band along posterior margin of segment (Figure 4 A); collophore and furcula white. Head. Antennae broken beyond second antennal segment. Eyes 8 (Figure 5 A), eyes G and H reduced; eye valley with five ciliate chaetae. Head with up to 11 Mc along antennal margin and four other Mc along inner margin of eye patch. Head dorsally (Figure 5 A) with seven anterior (A0, A2, A3, A5, S3, S4, S5) and one posterior (Pa5) Mc. Pre-labral chaetae finely denticulate, appearing smooth at low magnification. Labral chaetae 5, 5, 4, all smooth; chaeta length heterogeneous (Figure 5 D): proximal row (P) with middle chaeta (P0) longest and P1 shortest; medial row (M) with second chaeta (M1) longest and M2 shortest; distal row (D) with inner chaeta (D1) longer than outer chaeta (D2). Peristomal chaetae ciliate, bothriotricha-like. Proximal pleural chaeta (pps) denticulate, appearing smooth at low magnification, distal chaeta (dps) reduced, short and smooth (Figure 5 D). Basal chaeta of maxillary palp ostensibly shorter than distal chaeta (Figure 5 E); sublobal plate with two short chaetae. Maxillary head with three teeth and four lamellae; lamella 1 apically acuminate, with a subapical rounded expansion and short, straight papillae (Figure 5 B, C), lamella 2 apically rounded, densely packed with micropapillae (Figure 5 B). Lateral appendage of labial papilla E not quite reaching tip of papilla (Figure 5 F). Labial palp with five smooth proximal chaeta. Labial triangle with M1M2rEL1l2A1 ��� 5 (Figure 5 G); r smooth 3/4 as long as E; l2 short, spine-like. All post-labial chaetae coarsely ciliate; with six chaetae along cephalic groove. 1Species not included due to inadequate descriptions: C. camelinus Womersley, 1930 and C. shae ff eri B��rner, 1906. 2 1 + 1 means there are two columns, each with one seta; 2 + 2 means there are two columns each with two setae; 2 means there is only one column along the eye margin with two setae. Body. Body Mc 23/0241+7+0+5. Th. 2-Abd. 3 tergal S-chaetae as 11/011?3, S-microchaetae as 10/10100. Th. 2 dorsally flat; Th. 3 greatly developed (Figures 4 A, B and 5H), conic, extending posteriorly to cover medial section of Abd. 1 to intersegmental membrane with Abd. 2. Th. 2 with one S-chaeta and one S-microchaeta on lateroanterior corner, and two Mc in p3 complex (Figure 5 H); Th. 3 with one latero-anterior S-chaeta and three Mc in p2-3 complex. Abd. 1 with one anterior S-microchaeta; chaeta a6 absent. Abd. 2 (Figure 5 I) with two bothriotricha (m2, a5), two Mc (m3, m5), and S-chaeta as; number and distribution of lateral microchaetae unclear. Abd. 3 (Figure 6 A) with three bothriotricha (m2, a5, m5), four Mc (m3, am6, pm6, p6), S-chaeta as, S-microchaeta d2 inserted posterior to Mc p6, and five lateral mesochaetae. Abd. 4 (Figure 6 C) with four bothriotricha (B6, T2, T4, an secondary bothriotrix F3p?/F3 near latero-posterior margin); S-chaetae as and ps present, additional S-chaetae present but specimen condition does not allow us to ascertain their number. Pseudopore on Abd. 4 inserted in field just posterior to T4; tergum with eight inner Mc, one medial unpaired and seven paired (Figure 6 C): anterior triangle formed by paired A1 and one medial unpaired Mc; Mc A3 anterior and internal to pseudopore; Mc A5 and A6 near posterior margin of segment; Mc B3, B4 and B5 forming a group; lateral Mc E2, E3, E4, F1 and F3 present, E1 a microchaeta, E4 inserted anterior to D2, F2 absent. Posterior chaetae 23 ��� 34. Chaetotaxy of Abd. 5 as in C. sabanus (Figure 3 E). Legs. Trochanteral organ with up to 54 chaetae. Th. 3 claw complex as in Figure 6 B: tenent hair spatulate, relatively long, surpassing level of inner paired ungual teeth; claws with three inner teeth; basal teeth short, subequal, inserted on basal half of inner edge; unpaired tooth shorter than basal teeth, inserted on distal half of inner edge. Outer teeth ending on basal quarter of unguis (Figure 6 D); lateral teeth short but conspicuous; dorsal tooth short, clearly visible only on dorsal perspective. Unguiculus shorter on fore and middle legs than on hind legs, usually truncate, sometimes lanceolate, with a small inner tooth; posterior lamella smooth. Furcula. Dens with one inner and one outer row of spines. Mucro with five teeth, typical for genus, but unusually short, 0.6 ��� 0.8 as long as inner edge of hind claw (Figure 6 E). Remarks Campylothorax notidanus sp. nov. is the only member of the genus with a greatly enlarged, dorsal fin-shaped metathorax. The species is also unusual in having a greatly reduced dorsal chaetotaxy, with only three dorsal head Mc in row S, two Mc on Th. 2 p3 complex, three Mc on Th. 3 p2 ��� 3 complex and lacking Abd. 4 lateral Mc F2. The new species shares with C. hexosetosa sp. nov. the number and arrangement of inner Mc on Abd. 4, and the number and arrangement of lateral mesochaetae on Abd. 3, but otherwise it is quite different from all known Campylothorax (Table 2). The identity of thoracic Mc is unclear. The two Mc on Th. 2 are probably homologous with p2 and p3, whereas those on Th. 3 may be a2, p2 and a4. Unfortunately, all of the specimens studied are opaque and the microchaetae that could provide points of reference on the identity of the Mc are invisible., Published as part of Felipe N. Soto-Adames, 2016, Chaetotaxy of first-instar Campylothorax sabanus (Wray), and description of three new Campylothorax species from Hispaniola (Collembola, Paronellidae), pp. 1583-1612 in Journal of Natural History 50 (25) on pages 1593-1598, DOI: 10.1080/00222933.2016.1145272, http://zenodo.org/record/269880, {"references":["Mitra SK, Dallai R. 1980. Studies of the genus Campylothorax Schott, 1893 (Collembola Entomobryidae Paronellinae) with the description of a new species from Zaire. Monitore Zoologico Italiano, N. S. Supplement XII 9: 273 - 321.","Schott H 1927. Kamerunische Collembola. Meddelelser Linkopings Hogre Allmanna Laroverk Redogorelse 1 - 39 + 4 plates.","Gruia MM. 1983. Collemboles Arthropleones de Cuba recoltes par les expeditions cubano-roumaines en 1969 - 1973, II. Resultats des expeditions biospeleologiques cubano-roumaines a Cuba. 4: 191 - 205.","Schott H. 1893. Beitrage zur Kenntnis der Insektenfauna von Kameroon. I. Collembola. Bihang till Kongliga Svenska Vetenskaps-Akademien Handligar. 19: 1 - 28 + 7 plates.","Wray DL. 1953. New Collembola from Puerto Rico. J Agric Univ PR. 37: 140 - 150.","Womersley H. 1930. XXX. - On the Apterygota collected in British Guiana by the Oxford University expedition of 1929. Ann Mag Nat Hist Ser. 10: 305 - 317.","Borner C. 1906. Das System der Collembolen nebst Beschreibung neuer Collembolen des Hamburger Naturhistorischen Museums. Mitteilungen aus dem Naturhistorischen Museum. 23: 147 - 188."]}

Plutomurus eristoi sp. nov. Figs 25���37 Type locality. GEORGIA, Imereti region, Tskaltubo district, Dzedzileti village, Sataplia-Tskaltubo karst massif, Satevzia Cave, 42 �� 25 ' 52.01 "N, 42 �� 33 ' 58.12 "E, 273 m. alt. Type material. Holotype: Satevzia Cave (dark zone), 22.iii. 2012, female (code: CAU SATEVZIA 20120322 01). Paratypes (same data as Holotype): five specimens in slide [CAU SATEVZIA 20120322 0 2 to 06]; one specimen mounted on SEM stub [CAU SATEVZIA 20120322 07]. Deposited material: holotype and paratype 2 in MZNA; other paratypes in ISU. This new species was previously mentioned from Satevzia Cave as ��� Plutomurus sp. 7 ��� (Barjadze et al. 2015). Description. Body length up to 3.15 mm excluding antennae (holotype 2.9 mm) (mean 2.9 mm, n = 4). Body cuticular, colour brownish-grey, with patches of darker pigmentation over the whole body except on antennae, legs (trochanter to tibiotarsus) and dens. Scales cover antennal I and II segments, head, body and legs. Head. Eye patch small, eyes not seen with light microscopy; Ratio body length/antennae length 1.14 (n = 3) (holotype 0.98). Prelabral chaetae 3 + 3, labrum with 5, 5, 4 chaetae on tubercles and 4 curved hooks distally (Fig. 25); labial chaetae smooth with labial papillae as in Fig. 26. Maxillary palp trifurcate, distal papilla chaeta longer than basal chaeta; sublobal plate with four sublobals, all shorter than basal chaeta (Fig. 27). Body. Third leg with well developed trochanteral and femoral organs, with about 38 and 34 chaetae respectively, a few chaetae outstanding (Fig. 28). Ventral tube with 30���35 distal, 12 + 12 anterior and 80���100 posterior chaetae in one specimen. Legs scaled to tibiotarsus. Tibiotarsus III with one ventral spine-like chaeta, tibiotarsi I and II without spine-like chaetae (formula 001) (Fig. 29); claw long and narrow, with one small internal unpaired tooth (basal to the both internal lamellae, on which in some specimens there are two small teeth), at 20 % of distance from the claw base (Figs 30, 34, 34 ���); Emp. swollen basally, with 0���3 teeth in two internal lamellae starting from middle point and extending basally. Tenent hair acuminate, 3.3 times the length of claw III (3.5 in the holotype); ratio claw III, Emp. III and tenent hair 3.27: 1.75: 1 (n = 4); claw with a thin pseudonychium, 0.37 times length of inner edge of claw, and paired basal lateral teeth. Tenaculum with 4 + 4 teeth (Fig. 31); presence of chaetal socket not uncertain. Ratio of manubrium: dens: mucro 4.83: 7.67: 1 (n = 3); outer margin of dens with 3 thick, spine-like chaetae; dental formula variable: 5��� 7 I���II / 3��� 6 I 2��� 4 I 1��� 6 I to 4��� 5 I 1��� 4 I 1��� 2 I 1 I 1��� 3 I (the Arabic numbers means the number of small spines; dental large spines in Roman, bold and in italics) (Fig. 32); mucro with two basal and two distal teeth (Fig. 33). Chaetotaxy. Dorsal Mac on each side: Th. II without macrochaetae; Th. III with 1 macrochaeta (p 3); Abd. I���V with 3 (m 2, m 3, m 4), 3 (m 2, m 3, m 4), 4 (m 2, m 6, p 1, p 6), 1 (E 3) 3 (p 2, p 3, p 4) Mac, respectively. Bothriotrichal formula 0/ 2 ��� 1 /0��� 0���1 ��� 2 ���0; each bothriothrix bordered anteriorly with 1���7 small chaetae. Macrochaetotaxy and bothriotrichal patterns as in Figs 35���37. Discussion. Plutomurus eristoi sp. nov. is most similar to P. iwatensis Yoshii, 1991, P. kawasawai Yosii, 1956 and P. ortobalaganensis Jordana & Baquero, 2012 in Jordana et al. 2012 based on the following characters: absence of eyes, presence of 3 + 3 prelabral chaetae, presence of pointed tenent hair and absence of subapical teeth or denticles on mucro. All of these species are cave dwellers. The new species differs from P. i w a t e n s i s, P. kawasawai and P. ortobalaganensis in the shape of claw and Emp. Plutomurus eristoi sp. nov. has a long, narrow claw, and a basally swollen and apically distinctly pointed Emp., while the claw and Emp. shape of the three similar species mentioned above are typical for the genus Plutomurus, that is a wide claw, widest at base, and pointed Emp., not basally swollen. The new species differs from P. iwatensis by the number of distal teeth on mucro (two in the new species, four in P. i w a t e n s i s). P. eristoi sp. nov. differs from P. kawasawai in the number of spine-like chaetae on the hind tibiotarsus (1 vs. 2 in P. ka w as a w a i), and in the presence of empodial teeth in the new species and their absence in P. kawasawai. Plutomurus eristoi sp. nov. has 1 spine-like chaeta on the hind tibiotarsus, whereas P. ortobalaganensis has 2; and the new species lacks macrochaetae on Th. II and has one on Th. III, while Th. II and III have 6 and 2 macrochaetae, in P. ortobalaganensis respectively. Etymology. The species is dedicated to Dr. Eristo Kvavadze, who was supervisor of the first author. Ecology. Plutomurus eristoi sp. nov. is probably troglophilous species, as it possesses pigmented body and eye patches., Published as part of Barjadze, Shalva, Baquero, Enrique, Soto-Adames, Felipe N., Giordano, Rosanna & Jordana, Rafael, 2016, New diagnosis for species of Plutomurus Yosii (Collembola, Tomoceridae), with descriptions of two new species from Georgian caves, pp. 77-96 in Zootaxa 4126 (1) on page 87, DOI: 10.11646/zootaxa.4126.1.3, http://zenodo.org/record/262880, {"references":["Yoshii, R. (1991) A new species of Tomocerid Collembola from the Cave of Pref. Iwate. Annales of the Speleological Institute of Japan (Iwaizumi), 9, 1 - 2.","Yosii, R. (1956) Monographie zur Hohlencollembolen Japans. Contribution from the Biological Laboratory Kyoto University, 3, 1 - 109.","Jordana, R., Baquero, E., Reboleira, S. & Sendra, R. (2012) Reviews of the genera Schaefferia Absolon, 1900, Deuteraphorura Absolon, 1901, Plutomurus Yosii, 1956 and the Anurida Laboulbene, 1865 species group without eyes, with the description of four new species of cave springtails (Collembola) from Krubera-Voronya cave, Arabika Massif, Abkhazia. Terrestrial Arthropod Reviews, 5, 35 - 85. http: // dx. doi. org / 10.1163 / 187498312 X 622430"]}

Plutomurus revazi sp. nov. Figs 5–24, Table 2 Type locality. GEORGIA, Imereti region, Tskaltubo district, Kumistavi village, Sataplia-Tskaltubo karst massif, Satsurblia Cave, 42 ° 23 ' 16.59 "N, 42 ° 36 ' 22.35 "E, 290 m. alt. Type material. Holotype, female in slide: dark zone, 30.xi. 2014 (code CAU-SATSURBLIA 20141130 01). Paratypes (same data as holotype): three females in slide, dark zone, 30.xi. 2014 (CAU SATSURBLIA 20141130 0 3, 0 7, 09); one male in slide, dark zone, 30.xi. 2014 (CAU SATSURBLIA 20141130 04); two females in slide, twilight zone, 30.xi. 2014 (CAU SATSURBLIA 20141130 0 8 and 10); five females in slide, dark zone, 24.i. 2015 (CAU SATSURBLIA 20150124 0 2, 0 5, 0 6, 11, 12); one specimen mounted on SEM stub (CAU SATSURBLIA 20141130 13). Deposited material: holotype and paratypes 0 4, 0 6 and 0 8 in MZNA; remaining paratypes in ISU. Complementary material. Six specimens in slide (codes CAU PROMETHEUS- 20141130 0 1 to 06), and two specimens mounted on SEM stub (CAU PROMETHEUS 20141130 0 7 and 08), from GEORGIA, Imereti region, Tskaltubo district, Kumistavi village, Sataplia-Tskaltubo karst massif, Prometheus Cave, 42 ° 22 ' 38.05 " N, 42 ° 36 ' 3.40 " E, 175 m. alt., dark zone, 30.xi. 2014. Specimens 0 1 and 0 4 in MZNA, the rest in ISU. The new species was previously mentioned from Prometheus and Satsurblia caves as “ Plutomurus sp. 2 ” (Barjadze et al. 2015). Description. Body length up to 5.6 mm excluding antennae (holotype 3.6 mm) (Satsurblia Cave, mean 4.56 mm, n = 10; Prometheus Cave, mean 3.42 mm, n = 6). Body cuticular, colour brownish-grey, with patches of darker pigmentation over the whole body except on antennae, legs and dens; scales covering antennal I and II segments, head, body and legs; colour and scale patterns, and habitus as in Figs 5–6. Head. Eye patch small, eyes not visible under light microscope, while in SEM exterior faded 3-4 (5) cornea present. Ratio of body length to antennae length up to 1.31 (holotype 1.02) (Satsurblia Cave, mean 1.18, n = 6; Prometheus Cave, mean 0.99, n = 6). Prelabral chaetae 3 + 3; labrum with 5, 5, 4 chaetae arising from tubercles and with four curved hooks on distal part of labrum (Fig. 7). Maxillary palp trifurcate, chaeta on distal papilla longer than basal papilla; sublobal plate with four sublobals, all shorter than basal chaeta (Fig. 9). Labial papillae as in Fig. 8; proximal labial chaetae up to 16; labial chaetae smooth. Body. Third legs with well developed trochanteral and femoral organs, with about 17 and 40 chaetae respectively, a few chaetae longer than others (Fig. 10). Ventral tube with 65 distal, 49 + 49 anterior and 54 posterior chaetae in one specimen. Legs scaled to tibiotarsus. Tibiotarsus III with two ventral spine-like chaetae (Fig. 11), such chaetae absent on tibiotarsi I to II (formula 002). Claw on all legs with characteristic small, unpaired tooth between the two lamellae (Figs 12 –14, 18); number of internal unpaired teeth varying from 2 to 5, (Table 2), with a minute basal tooth at 20–40 % of the claw length from the base, at base of the two internal lamellae; three unpaired teeth usually present, second and third teeth occasionally absent, formed by the fusion of both lamellae; distal tooth formed by terminal fusion of lamellae (Fig. 18 ’); Emp. blade-like, usually with two internal serrated lamellae bearing 0–8 teeth starting near distal part and extending to base (Figs 12 –14, 18– 19; Table 2). Tenent hair long, clavate. Ratio of claw III: Emp. III: tenent hair 1.8: 1.1: 1 (holotype); claw with a thin pseudonychium, 0.76 times length of inner edge of claw and two basal and lateral teeth. Tenaculum with 4 + 4 teeth and one chaeta on corpus (Fig. 17). Ratio of manubrium: dens: mucro 5.7: 7.5: 1 (n = 8, including holotype). Outer margin of dens with four thick, spine-like chaetae; dental formula very variable (Table 2) (Fig. 16). Mucro with 2 basal, fused teeth, 0–2 intermediate denticles, and 2 distal teeth (Fig. 15). Male genital plate in Figs 20–21. Chaetotaxy. Dorsally Mac on each side: Th. II–III with 5 (m 3, m 4, m 5, p 3, p 4) and 2 (p 3, p 5) Mac, respectively; Abd. I–V with 3 (m 2, m 3, m 4) 3 (m 2, m 3, m 4) 4 (m 2, m 6, p 1, p 6) 3 (A 6, C 6, E 3) and 3 (p 2, p 3, p 4) Mac, respectively. Bothriotrichal formula 0/ 2 – 1 /0– 0–1 – 2 –0. Bothriothrix bordered anteriorly by 1–3 small chaetae. Macrochaetotaxy and Bothriotrichal pattern as in Figs 22–24. Specimen Claw I Emp. I Claw II Emp. II Claw III Emp. III dens dens apical number basal Holotype 3 + 1 b 3 + 3 2 + 1 b 4 + 3 3 + 1 b 5 + 4 7 III 7 I 2 I 3 I 3 + 1 b 5 +? 3 + 1 b 4 + 3 3 + 1 b 4 +0 10 II 4 I 3 I 2 I Paratype 1 3 + 1 b 2 +0 3 + 1 b - 3 + 1 b 5 + 3 23 II 4 I 7 I 2 I 1 I 0 I 3 + 1 b 5 + 2 3 + 1 b 7 + 5 - - 9 II 3 I 5 I 2 I 2 I Paratype 2 3 + 1 b 4 + 3 2 + 1 b 4 + 5 3 + 1 b 6 + 6 14 I 5 I 5 I 4 I 2 + 1 b 4 +? 2 + 1 b 4 +? 3 + 1 b - 17 III 4 I 5 I 1 I 4 I Paratype 3 3 + 1 b 4 + 5 3 + 1 b 3 + 1 3 + 1 b 6 + 3 15 III 2 I 2 I 3 I 4 I 3 + 1 b 2 + 3 - 4 + 4 3 + 1 b - - - Paratype 4 3 + 1 b 4 + 3 4 + 1 b 8 + 3 2 + 1 b 1 +0 - - 3 + 1 b 5 + 6 - - - - - - Paratype 5 3 + 1 b 2 +0 3 + 1 b - 3 + 1 b 4 + 5 14 II 6 I 4 I 3 I 1 + 1 b 3 +? 3 + 1 b 3 + 3 3 + 1 b 4 + 5 12 III 5 I 3 I 3 I 2 I Paratype 6 2 + 1 b 3 + 4 3 + 1 b 1 + 1 3 + 1 b 3 + 4 9 II 9 I 5 I 1 + 1 b 2 + 4 - - 3 + 1 b 4 + 4 12 IV 5 I 3 I 3 I Paratype 7 1 + 1 b - 2 + 1 b 4 + 5 1 + 1 b 7 + 7 11 II 6 I 4 I 3 I 2 + 1 b 4 + 3 1 + 1 b 7 + 3 2 + 1 b 3 + 4 12 II 3 I 6 I 2 I 3 I Paratype 8 2 + 1 b 3 + 2 - - 1 + 1 b 6 +? 12 II 9 I 2 I 3 I 2 + 1 b 3 + 2 - - - - 9 II 5 I 2 I 2 I 3 I Paratype 9 1 + 1 b 3 + 4 1 + 1 b 4 + 4? 1 + 1 b 6 + 5 11 II 8 I 4 I 3 I 1 + 1 b 4 +? 1 + 1 b 3 +? 1 + 1 b 6 + 4 15 II 8 I 2 I 4 I Paratype 10 3 + 1 b 4 + 3 4 + 1 b 4 + 4 3 + 1 b 6 +0 - - 3 + 1 b 2 + 2 4 + 1 b 2 + 2 3 + 1 b - - - Paratype 11 3 + 1 b 2 +0 2 + 1 b 4 +? - - 17 II 2 I 5 I 3 I 3 I 3 + 1 b 5 + 4 2 + 1 b 2 + 1 - - 17 II 6 I 3 I 4 I Range 1–3 + 1 b 2–5 + 0–6 1–4 + 1 b 1–8 + 1–5 1–3 + 1 b 1–7 + 0– 7 7–23 9–15 small small 1– 2 – 5 big spines 4 big spines Variable characters. The number of inner teeth on claw is variable. In the 12 specimens examined the number of inner claw teeth vary from 2 to 5; the number of teeth on inner two lamellae of the Emp. vary from 0 to 8. The number of dental spines is also variable. The number of small basal dental spines ranges from 7 to 23. In paratype 1 the dental spine formula on one dens is 23 II / 4 I 7 I 2 I 1 I 0 I, whereas on the other it is 9 II / 3 I 5 I 2 I 2 I. In most specimens the mucro lacks intermediate denticles, but in some specimens 1 or 2 intermediate denticles are present. Discussion. The new species is most similar to the North American P. grahami (Christiansen, 1980) based on the following characters: apparently blind under light microscopy, presence of 3 + 3 prelabral chaetae, and presence of clavate tenent hair. Both are cave-dwelling species. The new species differs from P. grahami by: (1) the formula of spine-like chaetae on the tibiotarsi: 0 0 2 in P. revazi sp. nov., while it is 0 0 1 in P. grahami; (2) dorsal macrochaetotaxy: 52 / 33433 in the new species, while it is 42 / 32413 in P. grahami. Etymology. The species is dedicated to Dr. Revaz Djanashvili, who, beginning in the 1960 s, was the first Georgian biospeleologist and collembolan taxonomist. Ecology. It is probably troglophilous species, because the body and eye patches are pigmented.

The Collembola fauna of the Galápagos Islands is relatively unexplored with only thirty-five reported species. Entomobryoidea, the most diverse superfamily of Collembola, is underrepresented, with only five species reported from the Galápagos. Here we present the findings of the first survey of Collembola from Galápagos lava tube caves, providing a significant update to the total number of entomobryoid Collembola species reported from the Galápagos Islands. Collections made during a March 2014 expedition to study lava tubes of the islands yielded new records for seven species of Entomobryoidea, including four genera not previously reported from the Galápagos Islands: Coecobrya, Entomobrya, Heteromurus, and Salina. As a result, three new species (Entomobrya darwini Katz, Soto-Adames & Taylor, sp. n., Pseudosinella vulcana Katz, Soto-Adames & Taylor, sp. n., and Pseudosinella stewartpecki Katz, Soto-Adames & Taylor, sp. n.) are described and new diagnoses are provided for Heteromurus (Heteromurtrella) nitens Yosii, 1964, Lepidocyrtus nigrosetosus Folsom, 1927 and Pseudosinella intermixta (Folsom, 1924). Lepidocyrtus leleupi Jacquemart, 1976 is synonymized with L. nigrosetosus. An updated checklist of all species within the superfamily Entomobryoidea reported from the Galápagos Islands is provided.

Key to Collembola species reported from Rapa Nui 1. Body globular, first four abdominal segments fused, antennae elbowed between segments 3 and 4....... Arrhopalites caecus - Body elongated, first four abdominal segments separated, antennae straight, not elbowed............................ 2 2. Third and fourth abdominal segments of approximately equal length; eyes absent; color white........... Folsomia candida - Fourth abdominal segment much longer than third segment; eyes present or absent; color variable...................... 3 3. Scales on head and body absent.......................................................................... 4 - Scales present........................................................................................ 7 4. Eyes and pigment present; mucro bidentate................................................................. 5 - Eyes and pigment absent; mucro falcate.................................................................... 6 5. Body pigment arranged as five longitudinal bands arranged dorsally and laterally; pigment on dorsum of fourth abdominal seg- ment arranged as a five-armed figure in adults, in juveniles as disconnected spots............ Entomobrya manuhoko n. sp. - Body pigment arranged as transverse bands on posterior body segments; pigment of dorsum of fourth abdominal segment appearing as a transverse zig-zag band................................................. Entomobrya multifasciata 6. Hind tibiotarsus with ventral row of seven smooth pointed setae............................ Coecobrya aitorerere n. sp. - Hind tibiotarsus with one ventral smooth, pointed seta......................................... Coecobrya kennethi 7. Mucro elongated, about half length of dens, with two strong teeth and minute third tooth; mucronal spine absent; dens with large scale-like setae............................................................. Cyphoderus manuneru n. sp. - Mucro short, less than one third length of dens, falcate or bidentate with subapical spine, dens only with small ventral scales.................................................................................................... 8 8. Mucro falcate; body with transverse bands of pigment........................................ Seira manukio n. sp. - Mucro bidentate, anteapical tooth present; pigment not in bands................................................. 9 9. Eyes 8 + 8, mesonotum projecting forward over thorax; body brown or orange....................... Lepidocyrtus olena - Eyes 4 + 4, mesonotum without forward projection; body blue........................... Pseudosinella hahoteana n. sp.

Trogolaphysa octosetosa sp. nov. Figs. 4 F, 18 A���F Table 4 Etymology. The epithet refers to the presence of 5 and 3 posterior macrochaetae on the meso- and metathorax, respectively. Material examined. Holotype female, in preparation. MEXICO, Yucatan, Chichen Itza, Berlese sample, leaf mould from moist woods and dry cenote, 25.vi. 1951, L. J. Stannard, coll., acc. no. 49763, INHS 579,835; 3 paratypes in preparation and 73 in alcohol (mostly small juveniles in poor condition) with same collection information as holotype. Description. Size up to 2.0 mm. Color pattern. Body with evenly distributed granular light blue pigment (Fig. 4 F), more intense on coxae; head lighter Head. Antennae up to 2.1 x head length. Guard sensillum of subapical sense organ relatively large, truncate; sensillar pit not seen. Sensilla in Ant. 3 sense organ laterally expanded. Eyes 6, eyes G and H absent; eye valley with 3 ciliate chaetae. Head dorsally with 2 anterior (A0, A 2) and 2 posterior (Pa 5, Pm 3) macrochaetae. Pre-labral chaetae weakly ciliate. Spines on distal margin of labrum distinct, separated. Labial triangle variable, M 1 and M 2 smooth or ciliate, either only one ciliate or both ciliate; r smooth and shortest of all; chaetae E, L 1 and L 2 ciliate; anterior chaetae smooth. Post-labial field with 6���9 ciliate chaetae between columns I and O. Body. Body macrochaetae 53 /0243+0+ 6. Mesothorax with 1 anterior (a 5) and 5 posterior (p 3 complex) macrochaetae (Fig. 18 A). Metathorax with 3 (a 2, p 2, p 3) macrochaetae (Fig. 18 B). Abd. 1 chaeta a 6 absent. Abd. 4 (Fig. 18 C) with pseudopore inserted in field posterior to T 4; macrochaetae A 3, A 5 and B 5 present; A 3 in field anterior to T 2; A 5 closer to pseudopore than B 5; 6 large lateral macrochaetae present. Posterior chaetae 9���10 + 9��� 10. Legs: Trochanteral with up to 28 chaetae. Metathoracic claw complex as in Fig. 18 D: tenent hair spatulate on all legs, 1.1 x as long as unguiculus; claws with 4 inner teeth, one basal tooth slightly wider and longer than other, proximal unpaired tooth shorter than basal teeth but well developed, distal unpaired tooth minute; teeth spread as 38 %, 42 %, 60 %, 86 %. Dorsal and lateral teeth ending on basal quarter of unguis, lateral teeth subequal to dorsal tooth. Unguiculus lanceolate, varying from narrow to normal, posterior edge smooth (Fig. 18 D). Furcula. Dens with 2 rows of ciliate spines: with up to 23 inner and 19 outer spines. Mucro with 4 sequential teeth (Figs. 18 E���F); mucro subrectangular in shape ��� 2.3 x as long as width of distal end of dens and 0.85 x as long as inner margin of metathoracic unguis. Remarks. Trogolaphysa octosetosa sp. nov. is the only Neotropical member of the genus with 6 eyes, 2 anterior dorsal head macrochaetae, 5 mesothoracic macrochaetae and 3 metathoracic macrochaetae. Six other species (T. carpenteri, T. cotopaxiana, T. palaciosi sp. nov., T. paracarpenteri sp. nov., T. relicta and T. subterranea) have 2 anterior head macrochaetae, but none has 3 metathoracic macrochaetae (Table 4). The color pattern of T. octosetosa sp. nov. is identical to that in T. paracarpenteri sp. nov., but in addition to the difference in number of thoracic macrochaetae, the species differ in the number (3 in T. paracarpenteri sp. nov., 4 in T. octosetosa sp. nov.) and shape of inner ungual teeth (basal teeth needle shaped in T. paracarpenteri sp. nov., keelshaped in T. octosetosa sp. nov.). The indistinct, diffuse blue color pattern of T. octosetosa sp. nov. is also seen in T. jataca, T. geminata, T. riopedrensis (Mari Mutt, 1987), T. subterranea, T. paracarpenteri sp. nov. and T. luquillensis (Mari Mutt, 1987), but the number of thoracic macrochaetae and details of unguis morphology distinguish all species. Trogolaphysa octosetosa sp. nov. was collected in sympatry with T. stannardi sp. nov. The paratypes of the two new species are mostly small juveniles, but color pattern and the relatively longer antennae in T. stannardi sp. nov. easily distinguish the species. In addition, adult T. stannardi sp. nov. are large and slender, whereas T. octosetosa sp. nov. adults are plump., Published as part of Soto-Adames, Felipe N., 2015, The dorsal chaetotaxy of first instar Trogolaphysa jataca, with description of twelve new species of Neotropical Trogolaphysa (Hexapoda: Collembola: Paronellidae), pp. 1-41 in Zootaxa 4032 (1) on pages 32-35, DOI: 10.11646/zootaxa.4032.1.1, http://zenodo.org/record/289744, {"references":["Mari Mutt, J. (1987) Puerto Rican species of Paronellidae (Insecta: Collembola). Caribbean Journal of Science, 23, 400 - 416."]}

Bernard, Ernest C., Soto-Adames, Felipe N., Wynne, J. Judson (2015): Collembola of Rapa Nui (Easter Island) with descriptions of five endemic cave-restricted species. Zootaxa 3949 (2): 239-267, DOI: http://dx.doi.org/10.11646/zootaxa.3949.2.6