Review of Lepidocyrtinus Börner, 1903 (Collembola, Entomobryidae, Seirinae): the African species Author Cipola, Nikolas Gioia Author Morais, José Wellington De 0000-0003-3801-7054 moraisjw@gmail.com Author Bellini, Bruno Cavalcante 0000-0001-7881-9436 entobellini@gmail.com text Zootaxa 2020 2020-12-24 4898 1 1 110 journal article 9066 10.11646/zootaxa.4898.1.1 9e4030d6-1154-422f-bb13-afd31181028f 1175-5326 4392515 3F66CE1D-1036-4226-BC28-029F9961E069 Lepidocyrtinus scheepersae sp. nov. Cipola & Bellini Figs 2 , 50–54 , Tables 1 and 4 Type Material. Holotype female on slide ( SAM / IZIKO : SAM-ENW-C009786): South Africa , Cape Town , Somerset West , Helderberg Nature Reserve , 34°03’40’’S , 18°52’20’’E ( Fig. 2 ), Southern Afrotemperate Forest , 142 m , 13.xi.2017 , beating and aspirating, A. Liu coll. 22 paratypes : 2 females on slides and 5 specimens in alcohol ( SAM / IZIKO : SAM-ENW-C009786); 2 females on slides ( SAM / IZIKO : SAM-ENW-C009786 donated to INPA- CLL 000033 ), same date as holotype ; 2 females and 1 juvenile in slides and 8 specimens in alcohol ( SAM / IZIKO : SAM-ENW-C009785), same data as holotype, except 17.x.2017 . Diagnosis. Body with dark bands on distal and proximal Ant III–IV and femur III distally ( Fig. 50 ); head mac A0 , A2–3 , A5 , M1–2 , M4i and S0–6 present; prelabral chaetae ciliated; labial papilla E with l.p. apically fingershaped and not reaching the base of a.a. ( Figs 51 C–D); basolateral and basomedian labial fields with chaetae R (subequal to others), E , L1–2 ciliated; Th II projected anteriorly and with 12–14 anterior, 8 median ( m1ip , m4–4p present, m4i mac absent) and 12–13 posterior mac ( p2ep and p3 as mac); Th III–Abd II with 11, 6, 4 central mac respectively; Abd IV with 16 central mac and 15–17 lateral mac ( Figs 52 , 53B ); unguis a.t. present; unguiculus pe lamella serrated and with a small proximal tooth; collophore anteriorly with 11 spine-like chaetae and 4 ciliated chaetae apically acuminate, posteriorly without spines, smooth chaeta present ( Fig. 54E ); manubrium ventrally with 2/2 subapical chaetae; manubrial plate with 3 blunt mac apically nipple-like; dens with 11–13 blunt mac apically nipple-like ( Figs 54 F–G). Description. Total length of body (head + trunk) 3.25–3.71 mm (n=4), holotype 3.71 mm . Specimens pale white with dark brownish to bluish pigments on distal and proximal Ant III–IV, distal Ant II, anterior and posterior regions of eyepatches, and dark blue pigments on coxa I and tibiotarsus I–III distally; eyepatches dark ( Fig. 50 ). Scales present on Ant I to proximal two thirds of Ant III, proximal one quarter of Ant IV, ventral and dorsal head, dorsal trunk, legs (except empodium), anterior collophore, ventral and dorsal manubrium and dens ventrally. Head. Antennae longer than the trunk length ( Fig. 50 ); ratio antennae: trunk = 1: 0.84–0.87 (n=4), holotype 1: 0.84; antennal segments ratio as I: II: III: IV = 1: 1.31–1.52: 2.29–2.74: 2.97–3.18, holotype 1: 1.35: 2.29: 2.97. Ant IV annulated on distal three quarters, with apical bulb apically unilobed. Ant III annulated on distal two thirds, apical organ with 2 elongated rods, 3 spiny guard sens, s-blunt sens of different sizes and ciliated chaetae. Ant I dorsally with 5 median mac and 3 smooth mic at the base, without spines ( Fig. 51A ). Eyes A and B larger, G and H slightly smaller, others subequal, and with 5 ‘IO’ chaetae; head dorsal macrochaetotaxy with 11–12 An , 4 A , 4 M and 8 S mac ( Fig. 51D ). Clypeal formula with 4 ( l1–2 ), 13–17 ( ft ), 3 ( pf0–1 ) ciliated chaetae, l1–2 and pf0–1 larger than others, l1 apically acuminate, 1–3 ft smaller, others subequal ( Fig. 51B ). Prelabral chaetae ciliated and not bifurcated. Labral p0 chaeta rarelly with minute median filament. Labral papillae with two inner conical projections, outer papillae absent. Maxillary palp with smooth t.a. and b.c. weakly ciliated, thicker and 1.08 longer than the t.a. Labial papilla E with l.p. apically finger-shaped and not reaching the base of a.a. ( Fig. 51C ). Basolateral and basomedian labial fields with chaetae M1–2 , R (subequal to others), E , L1–2 ciliated. Ventral head with about 29 ciliated chaetae, postlabial formula with 4 ( G1–4 ), 3 ( H2–4 ), 3 ( J1–3 ) chaetae, 13 thin posterior chaetae, and 1 larger b.c. surrounding the cephalic groove ( Fig. 51E ). FIGURE 50. Lepidocyrtinus scheepersae sp. nov. , habitus of a specimen fixed in alcohol (lateral view). Scale bars: 1 mm. Thorax chaetotaxy ( Fig. 52 A–B). Th II projected anteriorly, a , m and p series with 12–14, 8 and 12–13 mac, respectively. Th III a , m and p series with 4, 1 and 9 mac, respectively. Ratio Th II: III = 2.11–1.67: 1 (n=4), holotype 2.11: 1. TABLE 3. Comparison among Lepidocyrtinus species from continental Africa with 5 central mac on Abd I.

Characteristics	Lepidocyrtinus species
L. africanus sp. nov.	L. annulicornis 1, 4 (B̂rner, 1903)	L. botswanensis sp. nov.	L. capensis 2–4 Womersley, 1934	L. dayi ( Yosii, 1959 )	L. grisea 2,4 ( Womersley, 1934 ) comb. nov.	L. paucisensillum sp. nov.	L. ryozoi 3 nom. nov.
Colour pattern	head dark and spot on Abd III	all Ant I–IV and leg III	spots on coxae I–III	head and Th II anteriorly dark	body all blue	Head and trunk all dark blue	spots on joint of head and Th II	all Ant III–IV and femur I–III distaly
Ratio antennae: trunk	Ant </= body	Ant <body	Ant <body	Ant = body	Ant <body	Ant <body	Ant </= body	Ant </= body
Ant IV apical bulb	2	1*	1–2	2	2	1	?	?
Ant IV annulated	+	+	+	+	+	+*	?	+
Ant III annulated	+	+	+	+	+	?	+	+
Prelabral chaetae	ciliated	?	ciliated	?	smooth	?	ciliated	?
Head mac group	Pa3, Pm3, Pp3	–	?	–	?	+	?	–	?
Th II projected	+	+	+	+	–	–	+	+
Th II mac	m1–2 complex	6	6	6	6	6	6	5	5
m4i	–	?	–	?	+	?	–	?
PmA group	7	6	6	7	7–8	5–8	6	5
PmB group	3	3	3	3	3	3	3	3
PmC group	2	2	2	4	8–9	5	2	3
Th III central mac	9	9	9	8	11	11	8–9	9
Abd II central mac	4	4	4	4	3	4	3–4	4
Trochanteral organ	27	20	23	?	38	18	63	60
Unguis inner teeth	4	4	4	4	3	4	4	4
Unguiculus pe lamella	proximal tooth	–	–	+	+	+	–	+	–
margin	smooth	serrated	serrated	smooth	serrated	smooth	serrated	smooth
Manubrial plate	number	3	2–3	1	1–2	–	2–3?	3	3
modified mac	shaped	pointed	club	pointed	club	–	club	pointed	nipple
Dens modified mac	number	5–7	4*	3–4	2–3	1	2–3	9	9–10
shaped	bullet	club	bullet	club	nipple	club	nipple	nipple

Abbreviations: (*) doubtful characteristic; (+) present; (–) absent; (?) unknown. References: (1) B̂rner, 1903; (2) Womersley, 1934 ; (3) Yosii, 1959 ; (4) Coates, 1968 . FIGURE 51A–E. Lepidocyrtinus scheepersae sp. nov. : head; A , left Ant I chaetotaxy (dorsal side); B , clypeus chaetotaxy; C , labial papillae E (right side), arrow indicates l.p.; D , dorsal chaetotaxy (left side); E , complete postlabial chaetotaxy (right side). FIGURE 52A–D. Lepidocyrtinus scheepersae sp. nov. : dorsal chaetotaxy; A , Th II; B , Th III; C , Abd I; D , Abd II. FIGURE 53A–C. Lepidocyrtinus scheepersae sp. nov. : dorsal chaetotaxy; A , Abd III, B , Abd IV; C , Abd V. FIGURE 54A–G. Lepidocyrtinus scheepersae sp. nov. ; A–C , subcoxa I–III (outer side), respectively; D , trochanteral organ (inner side); E , collophore chaetotaxy (lateral view); F , manubrium ventral chaetotaxy; G , chaetotaxy of the manubrial plate and proximal dens (lateral view). Abdomen chaetotaxy ( Figs 52 C–D, 53A–C). Abd I a , m and p series with 1, 5 and 0 mac, respectively. Abd II a , m and p series with 1, 4 and 0 mac, respectively. Abd III a , m and p series with 1, 3 and 1 mac, respectively. Abd IV with 16 central mac on A– T series and 15–17 lateral mac on E– Fe series; at least 3 posterior sens ( ps type I, others type II ) and 8 posterior mes. Abd V a , m and p series with 1, 4 and 1/5 mac, respectively. Ratio Abd III : IV = 1: 3.86–4.62 (n= 4), holotype 1: 4.04. Legs. Subcoxa I with 6 chaetae on a row and 2 psp; subcoxa II with an anterior row of 13 chaetae, posterior row of 7 chaetae and 6 psp; subcoxa III with one row of 12 chaetae, 4 anterior chaetae and 2 posterior psp ( Fig. 54 A–C). Trochanteral organ with about 67 spine-like chaetae ( Fig. 54D ). Tibiotarsus not subdivided, outer side with 5 larger ciliated chaetae apically acuminate, inner side with 7 mac weakly ciliated. Unguis with 4 inner teeth, b.t. larger, m.t. subequal to a.t. in length. Unguiculus with all lamellae smooth and acuminate, except pe serrated and with a small proximal tooth; ratio unguis: unguiculus = 1: 0.54. Tibiotarsal smooth chaeta 1.79 larger than unguiculus and tenent hair 0.69 smaller than unguis outer edge. Collophore. Anterior side with 21–23 chaetae, 11 proximal spine-like chaetae, 1 thin and 3–5 ciliated chaetae, 4 ciliated chaetae apically acuminate and 2 mac distally; posterior side with 15 chaetae (2 unpaired), 14 thin ciliated chaetae widely distributed and 1 smooth chaeta distally; lateral flap with about 44 chaetae, 8 smooth and 36 ciliated ( Fig. 54E ). TABLE 4. Comparison among Lepidocyrtinus species from South Africa with 6 or 7 central mac on Abd I, and 4 central mac on Abd II.

Characteristics	Lepidocyrtinus species
L. barnardi ( Womersley, 1934 )	L. lindei 1 ( Coates, 1968 ) comb. nov.	L. marephila 1 ( Coates, 1968 ) comb. nov.	L. scheepersae sp. nov.
Colour pattern	spots on trunk and legs	body all dark blue	all Ant I and femur I–III	spots on femur II–III distally
Ratio antennae: trunk	Ant = body	Ant <body	Ant <body	Ant> body
Ant IV apical bulb	2	?	1*	1
Ant IV annulated	+	+	+	+
Ant III annulated	+	+	+	+
Prelabral chaetae	smooth	?	?	ciliated
Head mac	S4	+	?	–	+
Pp5	+	?	?	–
Th II projected	+	–	–	+
Th II mac	m1–2 complex	6	5	6	6
m4i	+	?	?	–
PmA group	7	7	7	6–7
PmB group	3	3	3	3
PmC group	5–6	6	5	3
Th III central mac	10	12	12	11
Abd I central mac	6	7	6	6
Trochanteral organ	94	?	?	67
Unguis inner teeth	3–4	4	4	4
Unguiculus pe lamella	proximal tooth	+	–	–	+
margin	serrated	smooth	serrated	serrated
Manubrial plate	number	–	2–3	2–3	3
modified mac	shaped	–	club	club	nipple
Dens modified mac	13–19	?	3	11–13

Abbreviations: (*) doubtful characteristic; (+) present; (–) absent; (?) unknown. References: (1) Coates, 1968 . Furcula. Manubrium ventral formula with 0, 0, 0, 2/2 (subapical), 20–22 (apical) ciliated chaetae of different sizes plus approximately 15 elongated apical scales per side ( Fig. 54F ); manubrial plate with 3 blunt mac finely ciliated and apically nipple-like, 14 ciliated chaetae of different sizes and 3 psp. Dens dorsally with one proximal row of 11–13 blunt mac finely ciliated and apically nipple-like, holotype with 11–13 ( Fig. 54G ) Etymology. The new species is named after the Collembola researcher Dr. Charlene Janion-Scheepers (SAM/ IZIKO, South Africa ), who provided us specimens of the new species. Remarks. Considering the continental taxa from Africa, L. scheepersae sp. nov. resembles L. barnardi and L. marephila ( Coates, 1968 ) comb. nov. in Th II with 6 median mac in m1–2 complex and Abd I–II with 6 and 4 central mac, respectively ( Table 4 ). Lepidocyrtinus scheepersae differs from these species by head mac Pp5 absent (present in L. barnardi ), Th II projected anteriorly (normal in L. barnardi and L. marephila ) and with 12–13 posterior mac ( 15–16 in L. barnardi , 15 in L. marephila ). In Th III it differs by 11 central mac ( 10 in L. barnardi , 12 in L. marephila ), and Abd IV with 16 central mac ( 10–11 in L. barnardi ). It also differs in ventral manubrium only with ciliated chaetae (3 distal smooth chaetae in L. marephila ) and dens with 11–13 blunt chaetae ( 2–3 in L. marephila , 13–19 in L. barnardi ). Lepidocyrtinus scheepersae also resembles L. betamponensis sp. nov. from Madagascar in habitus, color pattern with distal spots of pigment on femur III, overall chaetotaxy pattern and other general morphology ( Figs 50–54 , 72–76 ). However, L. scheepersae differs from this species by dorsal head mac S4 present (absent in L. betamponensis ), prelabral chaetae ciliated and basomedian labial fields with chaetae E ciliated (both smooth in L. betamponensis ). It differs also by dorsal chaetotaxy of the Th III–Abd II with 11, 6, 4 central mac, respectively (8–10, 1–2, 3– 4 in L. betamponensis ) and Abd IV with 16 central mac ( 12 in L. betamponensis ). Other differences between them are: collophore anteriorly with 11 spine-like chaetae but absent on posterior side in the new species (9 anterior and 8 posterior in L. betamponensis ) and manubrial plate and dens with blunt mac apically nipple-like (apically bullet-like and abruptly acuminate at the apex in L. betamponensis ). Key to continental African species of Lepidocyrtinus 1 Abd I with 3 or less central mac ( Fig. 41C )................................................................. 2 - Abd I with 5 or more central mac ( Fig. 11A , 15C )........................................................... 9 2 Th III with 3 central; Abd I without mac................................................................... 3 - Th III with 4 or more central mac; Abd I with at least 1 central mac ( Fig. 41 B–C).................................. 4 3 Th II with 3 central mac in a triangle shape; Th II normal; unguis a.t. present.......... L. metala ( Coates, 1968 ) comb. nov. - Th II with 3 central mac in a linear shape; Th II projected anteriorly; unguis a.t. absent. L. tsikama ( Coates, 1968 ) comb. nov. 4 Abd I–II with 1 and 3 central mac, respectively ( Figs 41 C–D).................................................. 5 - Abd I with 2–3 central mac, Abd II with 2 or 4 central mac ( Fig. 35D )........................................... 6 5 Th II with 4–5 posterior mac ( Fig. 41A ); Th III with 4 central mac ( Fig. 41B ); unguiculus pe lamella serrated................................................................................................... L. fabioi sp. nov. - Th II with 8 posterior mac; Th III with 6 central mac; unguiculus pe lamella smooth but with one proximal tooth.................................................................................. L. rykei ( Coates, 1968 ) comb. nov. 6 Abd I with 2 central mac............................................................................... 7 - Abd I with 3 central mac............................................................................... 8 7 Th II with 4 mac in m1–2 complex and 8 posterior mac; Th III with 4 central mac (as Fig. 41B ); Abd II with 4 central mac (as Fig. 35D )................................................................ L. annela ( Coates, 1968 ) comb. nov. - Th II with with 5 mac in m1–2 complex 10 posterior mac; Th III with 8 central mac; Abd II with 2 central mac................................................................................ L. mathewsi ( Coates, 1968 ) comb. nov. 8 Th II with 10 posterior mac (PmA–C group); Th III with 9 central mac, p1i and p3 mac present; dens dorsally with 4 modified mac................................................................ L. metarsiosa ( Coates, 1968 ) comb. nov. - Th II with 9 posterior mac (PmA–C group); Th III with 7 central mac, p1i and p3 mac absent; dens dorsally with 2–3 modified mac................................................................. L. vaneedeni ( Coates, 1968 ) comb. nov. 9 Abd I with 6–7 central mac ( Fig. 52C ).................................................................... 10 - Abd I with 5 central mac ( Fig. 11A )..................................................................... 13 10 Th II with 6 median mac in m1–2 complex ( Fig. 15A ); Abd I with 6 central mac ( Fig. 52C ); unguiculus pe lamella serrated (as Fig. 37F )........................................................................................... 11 - Th II with 5 median mac in m1–2 complex (as in Fig. 12A ); Abd I with 7 central mac; unguiculus pe lamella smooth (as Fig. 12C )................................................................... L. lindei ( Coates, 1968 ) comb. nov. 11 Head with S4 mac ( Fig. 14F ); Th III with 11 or less central mac ( Fig. 52B ); dens with 10 or more modified mac ( Fig. 17G ).. .................................................................................................. 12 - Head devoid of S4 mac (as Fig. 19E ); Th III with 12 central mac; dens with 2–3 modified mac............................................................................................. L. marephila ( Coates, 1968 ) comb. nov. 12 Prelabral chaetae smooth ( Fig. 14D ); head with Pp5 mac ( Fig. 14F ); Abd IV with 10–11 central mac ( Fig. 16B ); manubrial plate without modified mac ( Fig. 17G )................................................. L. barnardi ( Womersley, 1934 ) - Prelabral chaetae ciliated; head without Pp5 mac ( Fig. 51D ); Abd IV with 16 central mac ( Fig. 53B ); manubrial plate with 3 modified mac ( Fig. 54G )............................................................. L. scheepersae sp. nov. 13 Th II with 5 median mac in m1–2 complex ( Figs 12A , 47A ).................................................. 14 - Th II with 6 median mac in m1–2 complex ( Fig. 35A )....................................................... 15 14 Th II PmA and PmC groups with 6 and 2 mac ( p3 as mic), respectively; Th III with p1p2 mac absent ( Fig. 47 A–B); unguiculus pe lamella serrated (as in Fig. 37F )................................................... L. paucisensillum sp. nov. - Th II PmA and PmC groups with 5 and 3 mac ( p3 as mac), respectively; Th III with p1p2 mac ( Fig. 12A ); unguiculus pe lamella smooth ( Fig. 12C )................................................................. L. ryozoi nom. nov. 15 Th III with 11 central mac ( Fig. 20B ).................................................................... 16 - Th III with 9 or less central mac ( Fig. 25B )................................................................ 17 16 Th II with 18–20 posterior mac ( Fig. 20A ); unguis a.t. absent and unguiculus pe lamella serrated; manubrial plate devoid of modified mac ( Fig. 22G )................................................................ L. dayi ( Yosii, 1959 ) - Th II with 16 or less posterior mac; unguis a.t. present and unguiculus pe lamella smooth (as in Fig. 12C ); manubrial plate with 2–3 modified mac...................................................... L. grisea ( Womersley, 1934 ) comb. nov. 17 Head dark and trunk with a pattern of spots of pigment (e.g. Fig. 23A ; Womersley 1934 , pag. 462); Th II PmA group with 7 mac............................................................................................... 18 - Head pale and trunk without a clear pattern of spots of pigment (e.g. Fig. 32 ); Th II PmA group with 6 mac............. 19 18 Th II PmC group with 2 mac; Th III with 9 central mac ( Figs 25 A–B); manubrial plate and dens with 3 and 5–7 modified mac, respectively ( Fig. 29C ).................................................................. L. africanus sp. nov. - Th II PmC group with 4 mac; Th III with 8 central mac; manubrial plate and dens with 1–2 and 2–3 modified mac, respectively......................................................................... L. capensis Womersley, 1934 19 Femur and tibiotarsus III distally with dark violet pigments; manubrial plate with 2–3 modified mac ( Fig. 11B )........................................................................................ L. annulicornis (B̂rner, 1903) - Legs pale ( Fig. 32 ); manubrial plate with 1 modified mac ( Fig. 38C ).......................... L. botswanensis sp. nov. Remarks. The species as L. armillatus ( Wahlgren, 1908 ) and L. laetus (B̂rner, 1908), L. annulipes Handschin, 1929 and L. flavovirens (B̂rner, 1903), are herein transferred to Lepidocyrtinus due to the presence of Ant III–IV annulated, unguis with a pair of large laterobasal teeth, dens dorsally with modified mac (at least in the first two species), and overall macrochaetotaxy pattern (known only to the last two species) (see B̂rner 1903; Wahlgren 1908 ; Denis 1924 ; Handschin 1929 ; Yosii 1959 ; Coates 1968 ). However, such species are not included in the key because they are in need to be redescribed, since previous descriptions are insufficient to establish specific diagnoses to them. In this sense, some Seira species from Africa which have Ant III and IV annulated and large laterobasal teeth on unguis, but also have poor descriptions unable to clarify their position among the Seirinae , such as S. addoensis Coates, 1968 , S. eleana Coates, 1968 , S. ethiopica ( Denis, 1924 ) and S. laeta (B̂rner, 1908) may also represent Lepidocyrtinus taxa as well. For this dubious condition, they are herein maintained as Seira s. lat. , as well as other species from India described in Lepidocyrtinus by Paliwal & Baijal (1985) . Madagascar species