Abstract
Herein we describe two new species of Brazilian springtails, both from Rio Grande do Norte, Brazil. Brachystomella nordestina sp.n. resembles other Neotropical congeners with 8+8 eyes, postantennal organ with four lobes and 19/19/18 chaetae on tibiotarsi I–III, but it is unique in the combination of trilobed apical bulb, 2+2 dorsal chaetae on thorax I, male’s genital plate with 4+4 modified eugenital chaetae and 2+2 hr chaetae on lateral anal valves. Seira (Lepidocyrtinus) dapeste sp.n. is similar to other Neotropical species of Lepidocyrtinus Börner, 1903 in long antennae, mesonotum projected over head and presence of modified blunt macrochaetae on dorsal furca, but differs in the combination of dorsal head chaetotaxy, 14–19 a macrochaetae on thorax II, 7–9 central macrochaetae on thorax III and 4 central macrochaetae on abdomen II. We also present a detailed comparison and an updated key of the Neotropical species of Brachystomella Ågren, 1903.
ZooBank: http://zoobank.org/943CFF4B-BC5E-42A9-93AA-7734A52126FB
Introduction
Brachystomella Ågren, 1903 is the largest genus of Brachystomellidae Stach, 1949 (Poduromorpha), and currently includes almost 70 valid species, nine of which are known from Brazil (Bellinger et al. 1996–2017; Abrantes et al. 2010, 2012). It is cosmopolitan, but most species are known from Neotropical Region (Najt & Weiner 1996, Weiner & Najt 2001). The genus is recognized from other Brachystomellidae especially by the following combination of characters: antennomere IV apical bulb simple to trilobed, eyes and postantennal organ present with vesicles in one line, mandibles completely absent, maxilla capitulum globular with rostral teeth, furca present, and anal spines absent (a complete diagnosis to the genus is presented in Systematics section) (Jordana et al. 1997).
Seira Lubbock, 1869 is the fourth largest genus of Entomobryidae Tömösvary, 1882 with 205 described species worldwide, about 25% of which are recorded from Neotropical Region (Bellinger et al. 1996–2017). The genus was divided in three subgenera by Yosii (1959): Lepidocyrtinus Börner, 1903, with modified blunt macrochaetae on dorsal furca; Afroseira Yosii, 1959 to include the single species S. rowanorum Yosii, 1959, with 2+2 bothriotricha on abdominal segment IV; and Seira s. str. Lubbock, 1869 to include the species without blunt macrochaetae on furca and 3+3 bothriotricha on abdominal segment IV. Since most species of the genus fit Seira s. str., this division of subgenera was mostly omitted in recent species descriptions (as in Cipola et al. 2014a, b; Godeiro & Bellini 2014, 2015). However, Neotropical adult specimens of Lepidocyrtinus can be easily separated from Seira s. str. taxa due the presence of modified blunt macrochaetae on furca, overall larger body size, long antennae, mesonotum strongly projected over head, and at least to some species, the presence of several anterior macrochaetae (excluding anterior collar) on mesonotum (Arlé 1959; Godeiro & Bellini 2014, 2015). All Neotropical species which fit the Lepidocyrtinus diagnosis are endemic to Brazil.
Herein we describe two new species of Brazilian springtails: Brachystomella nordestina sp.n. and Seira (Lepidocyrtinus) dapeste sp.n., both from Natal, Rio Grande do Norte State. We present a comparison among Neotropical species of Brachystomella and update the key to those species presented in Weiner & Najt (2001) and also provide some comments on Neotropical Lepidocyrtinus.
Material and methods
Specimens were collected with entomological aspirators and pitfall traps, preserved in 70% ethanol and posteriorly cleared with Nesbitt’s solution and mounted in microscopic slides in Hoyer’s solution. Drawings and measurements were made based on all type series, with a microscope using a drawing tube. Photographs were made using an optical microscope (Nikon Eclipse NiU) attached to a digital camera (Nikon DS-Ri1).
Terminology and chaetotaxy schemes used in the description of Brachystomella nordestina sp.n. followed mainly: Yosii (1960) with additions of Christiansen & Bellinger (1980) to dorsal chaetotaxy, and as presented in Jordana et al. (1997), Fjellberg (1998) and Thibaud et al. (2004); D’Haese (2003) to dorsal sensilla on antennomere IV; Cipola et al. (2014b) to labral chaetotaxy; Fjellberg (1999) to labial palp structures, except to proximal chaetae of labial palp, which followed D’Haese (2003); Massoud (1967) to labial basomedian and basolateral fields chaetae; and Deharveng (1983) as presented in D’Haese (2003) to tibiotarsal chaetae. Description of Seira (Lepidocyrtinus) dapeste sp.n. followed: Mari Mutt (1979) with modifications of Soto-Adames (2008) to dorsal head chaetotaxy; Szeptycki (1979) and Soto-Adames (2008) to dorsal tergal chaetotaxy; Zhang & Deharveng (2015) to specialized chaetae (sensilla and microsensilla); Cipola et al. (2014b) to labral chaetotaxy; Fjellberg (1999) to labial palp structures; Gisin’s system (1967) with addictions of Mari Mutt (1986a) to labial chaetotaxy (all chaetae presented in uppercase). Abbreviations used in the descriptions are: Ant. = antennal segment; Th. = thoracic segment; Abd. = abdominal segment; PAO = postantennal organ; ms = microsensillum; subapical organite = or.
Type materials of the species were deposited at the Collembola Collection of Centro de Biociências of Universidade do Rio Grande do Norte, Natal, Rio Grande do Norte, Brazil (CC/UFRN) and at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland (ISEA).
Results
Taxonomic summary
Brachystomellidae Stach, 1949
Brachystomella Ågren, 1903
Type species: Brachystomella parvula (Schäffer, 1896)
Genus Diagnosis
Specimens color varying from dark blue to colorless, tegument weakly to heavily granulate. Body chaetae smooth to serrate. Ant. IV with eversible apical vesicle simple, bi or trilobed; truncate subapical organite and dorso-lateral sensillum present; 5 to 8 dorsal sensilla present, sometimes weakly differentiated from ordinary chaetae; ventral side usually with some medium sized blunt to truncate chaetae. Ant. III sense organ typically with 5 elements: 2 small sensory rods partially covered by basal tegument fold, 2 guard sensilla of subequal to clearly different sizes, and 1 ventrolateral ms. Ant. I and II with 7 and 10–13 chaetae respectively. Oral cone short and triangular. Mandibles totally absent; maxillae with cardo, globular capitulum with up to 10 rostral teeth. Labial palps and maxillary outer lobes reduced. Labral region with 2334 chaetae. Pre-labral central region with 2 chaetae. PAO rounded with 3 to 10 vesicles in one single row. Eyes 8+8 to 2+2. Thoracic and abdominal segments clearly divided. Th. I with 2+2 to 4+4 chaetae. Th. I –III microsensilla formula 0/1/0. Trunk long sensorial chaetae half tergum formula from Th. II to Abd. V as: 22/11111, 22/21111 or 22/22211. Ungues with a pair of lateral teeth, unguiculi usually absent. Ventral tube with 3+3 chaetae. Tenaculum usually present; corpus lacking chaetae, each ramus with 1 to 3 teeth. Furca present with three different morphologies: well developed, with 5 to 6 (rarely 7) dorsal chaetae on dens and long mucro; reduced, with 3 (rarely 4) dorsal chaetae on dens and short mucro; or vestigial, with dens reduced to stubs lacking chaetae and mucro. Males with or without modified eugenital chaetae (secondary sexual characters). Lateral anal valves usually with some reduced chaetae (hr). Anal spines absent (adapted after Jordana et al. 1997; Weiner & Najt 2001; Queiroz & Weiner 2011).
Brachystomella nordestina sp.n. Souza, Bellini & Weiner
Habitus of Brachystomella nordestina sp.n. (specimen fixed in ethanol, lateral view).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Brachystomella nordestina sp.n. antennae: 2, Ant. IV and III dorsal view (captions to part of dorsal sensilla and i chaeta on Ant. IV); 3, Ant. IV ventral view (arrows point to some ventral blunt chaetae); 4, Ant. IV and III dorsal view; 5, Ant. IV and III ventral view.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Brachystomella nordestina sp.n.: 6, Pre-labral and labral chaetotaxy; 7, right maxilla capitulum; 8, left eye patch and PAO; 9, complete labial chaetotaxy; 10, left maxillary outer lobe; 11, ventral head chaetotaxy (left side); 12, dorsal body chaetotaxy (arrows point to chaetae present or absent).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Brachystomella nordestina sp.n.: 13, leg III (posterior view); 14, tibiotarsus I schematic chaetotaxy (posterior view); 15, tibiotarsus III and unguis (posterior view); 16 tibiotarsus III schematic chaetotaxy (posterior view); 17, ventral tube (ventral view); 18, tenaculum rami (ventral view); 19, right dens and mucro; 20, female genital plate and surrounding chaetae (arrows point to chaetae present on adults); 21, male genital plate and surrounding chaetae; 22, anal valves chaetotaxy (ventral view, arrows point to chaetae present or absent).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
ZooBank: http://zoobank.org/85E83705-8AB9-4339-9163-5E971E488C94
Type material
Holotype female, Brazil, Rio Grande do Norte State, Natal Municipality, Mata do Sagui, Campus Central da Universidade Federal do Rio Grande do Norte (5°50’34.30”S 35°12’04.63”W), 08-ix-2015, N. Santos, P. Souza & M. Freire coll., on slide without number, deposited in CC/UFRN. Paratypes: 4 females, same data as holotype; 3 males (1 subadult) and 11 females, same data as holotype, except collection date (11-iii-2015); 1 male and 9 females (one subadult), same data as holotype, except collection date (03-vi-2015), all on slides without number, deposited in CC/UFRN. Other paratypes: 1 male and 6 females, same data as holotype, deposited in ISEA.
Diagnosis
Specimens pigmented, body chaetae smooth. Ant. IV with trilobed apical bulb, with 6 dorsal subcylindrical sensilla and 10 ventral modified blunt chaetae. Ant. III sensory organ guard sensilla of subequal sizes. Eyes 8+8. PAO with 4 lobes (rarely with 3). Dorsal head with chaetae a0, sd1 and c1–2 present. Trunk dorsal ordinary chaetae short; long sensory chaetae smooth and acuminate with half tergum formula from Th. II to Abd. V as: 22/21111. Th. I with 2+2 dorsal chaetae. Subcoxa 1 of I leg with 1 chaeta. Tibiotarsi I–III with 19, 19 and 18 acuminate chateae, including tenent-hairs. Unguiculus absent. Tenaculum with 3+3 teeth. Furca well developed with 5 dorsal chaetae on each dens, mucro straight with apex slightly hooked. Male genital plate with 4+4 modified eugenital strong chaetae (secondary sexual character). Lateral anal valves with 2+2 hr microchaetae.
Description
Body bluish, purple or brownish (Fig. 1). Body granules medium sized, uniformly distributed on tegument. Body (head + trunk) length of holotype 1074 μm, paratypes maximum body length, 1430 μm, minimum 570 μm, average 1120 μm. Body chaetae smooth.
Head. Antennae slightly longer (274 μm in holotype) than head length (264 μm in holotype). Ant. IV with trilobed apical bulb, dorsally with 6 subcylindrical sensilla (S1,S3–4,S7–9), reduced chaeta i, dorsolateral ms and truncate or; ventrally with 10 modified blunt chaetae (Figs. 2–5). Ant. III sense organ with 2 small sensory rods partially covered by basal tegument fold, 2 guard sensilla of subequal sizes and 1 ventrolateral ms (Figs. 4–5). Ant. IV and III fused dorsally, ventral separation well marked (Figs. 4–5). Ant. I and II with 7 and 12 chaetae, respectively. Prelabral region with 1+1 central and 1+1 lateral chaetae (from clypeus) (Fig. 6). Labral region with 2(pp1) / 3(p0–1), 3(m0–1), 4 (a1–2) chaetae (Fig. 6). Maxillae with 6–8 rostral teeth (Fig. 7). Eyes 8+8 with 3 or 2 interocular chaetae (oc1–3, oc1 present or absent); PAO with 4 lobes (abnormally in one specimen with 3 on one side of head), about 1.8 times the size of ocular lens B (Figs. 8 and 12). Dorsal head chaetotaxy as in Fig. 12, with short ordinary chaetae; chaetae a0, sd1 and c2 present; sd2, sd4, oc1 and c5 present or absent. Labium with one distinct papilla and 4 proximal chaetae (a1–2, p1–2); basomedial field with 4 chaetae (E, F, G, f); basolateral field with 4 chaetae (e’, d, c, b), e chaeta present (Fig. 9, e chaeta not shown). Maxillary outer lobe partially fused with anterior region of labial basolateral field, with two chaetae (none of them apical) (Fig. 10); sublobal plate absent. Postlabial chaetotaxy as in Fig. 11, 2+2 chaetae surrounding the ventral groove.
Trunk main chaetotaxy. Trunk dorsal chaetotaxy as in Fig. 12. Ordinary chaetae short; long sensory chaetae (s) smooth and acuminate with half tergum formula from Th. II to Abd. V as: 22/21111. Th. I with 2+2 dorsal chaetae. Th. II–III dorsal chaetotaxy similar, with 3+3 chaetae on ‘a’ series (a1, a4 and a6), 2+2 on ‘m’ series (m5 and m7, m5 can be absent on Th. III – seen in one specimen) and 5+5 on ‘p’ series (p1–2, p4–6); Th. II long sensory chaetae as p4 and m7; lateral ms present on Th. II. Thoracic sterna without chaetae. Abd. I dorsal chaetotaxy with p5 (p4 in Weiner & Najt 2001) and p6 as long sensory chaetae. Abd II–IV only with p5 as long sensory chaeta; Abd. V with p2 as long sensory chaeta. Abdominal sternum I without chaetae, sternum II with 1+1 chaetae.
Legs. Subcoxae 1 of I–III legs with 1, 2, 2 chaetae; subcoxae 2 I–III with 0, 2, 2 chaetae; coxae I–III with 3, 5, 7 chaetae; trochanters I–III with 4–5, 5, 4 chaetae; femora I–III with 12, 11, 10 chaetae; tibiotarsi I–III with 19, 19 and 18 acuminate chateae, respectively; A1 as long acuminate tenent-hair, M present, B7 absent on tibiotarsus III (Figs. 13–16). Anterior and posterior pretarsal chaetae present (Fig. 15). Ungues with one inner tooth at proximal 1/3 the length of its inner edge, a pair of lateral teeth and one discrete outer tooth (Fig. 15). Unguiculi absent.
Abdominal appendages. Ventral tube short, with 3+3 chaetae (Fig. 17). Tenaculum corpus without chaetae, each ramus with 3 teeth (Fig. 18). Furca well developed with 5 dorsal chaetae on each dens (99 μm in holotype); mucro (37 μm in holotype) straight with apex slightly hooked (Fig. 19); ratio mucro : dens = 1 : 2.67.
Genital plates and anal valves. Female genital plate as in Fig. 20, with 2+2 pregenital, 7–9 circumgenital and 0–2 eugenital chaetae. Male genital plate as in Fig. 21, with 3+3 pregenital, 11 circumgenital and 4+4 modified eugenital strong chaetae (secondary sexual characters). Lateral (even) anal valves with 13–14 chaetae on each valve plus 2+2 hr microchaetae (3+3 abnormally only in holotype) (Fig. 22). Dorsal anal valve with 1+1 chaetae plus 1+1 hr microchaetae (Fig. 22).
Etymology
“nordestina” is a Portuguese word which refers to someone “who was born in northeastern region” of Brazil. The word carries stigma since a large amount of poor people live in this region. We name this species as Brachystomella nordestina sp.n. to honor this group of Brazilians.
Habitat
Specimens of Brachystomella nordestina sp.n. were collected from leaf litter over sandy soil of an urban forested area surrounding Federal University of Rio Grande do Norte campus (Natal, Rio Grande do Norte State, northeastern region of Brazil), which is part of the Atlantic Forest phytogeographic domain. Specimens were found both during rainy and dry seasons. Climate of the area is ‘As’ following the Köppen-Geiger system, which means an equatorial savannah-like hot climate with a dry summer (Kottek et al. 2006).
Remarks
Brachystomella nordestina sp.n. resembles other Neotropical species of the genus with trilobed apical bulb, 8+8 eyes, PAO with four lobes, chaetae smooth and 19/19/18 chaetae on tibiotarsi I–III such as B. purma Weiner & Najt, 2001 and B. victoriensis Izarra, 1972. However, the new species differs from B. purma in 2+2 hr chaetae on lateral anal valves (1+1 in B. purma); and from B. victoriensis in Th. I with 2+2 chaetae (3+3 in B. victoriensis), absence of capitate chaetae on tibiotarsi (2 in B. victoriensis) and 5 dorsal chaetae on dens (6 on B. victoriensis). The new species is also similar to the widespread B. agrosa Wray, 1953 (see Table 1), but it differs in Ant. IV apical bulb trilobed (unilobed in B. agrosa) and male’s genital papillae with 4+4 modified eugenital chaetae (3+3 in B. agrosa). A detailed comparison among the Neotropical species of Brachystomella is present in Table 1.
Key to Neotropical species of Brachystomella (updated from Weiner & Najt, 2001)*
- 1. Less than 8+8 eyes present2
- 1’. 8+8 eyes present7
- 2 (1). 7+7 to 5+5 eyes present3
- 2’. 2+2 eyes present, thoracic tergum I with 2+2 chaetae, Ant. IV apical bulb trilobed, dens with 5 chaetae eachgarayae Queiroz & Weiner, 2011; Brazil
- 3 (2). 7+7 or 6+6 eyes present4
- 3’. 5+5 eyes present5
- 4 (3). 7+7 eyes present, PAO with 5 vesicles, dens each with 5 chaetae septemoculata Denis, 1931; Costa Rica, Jamaica, the French West Indies, Cuba, Mexico
- 4’. 6+6 eyes present, PAO with 5–9 vesicles, dens each with 6 chaetae sexoculata Massoud, 1967; Peru
- 5 (3’). Thoracic tergum I with 2+2 chaetae, PAO with 5 vesicles, even anal valves each with 1 or 2 chaetae hr 6
- 5’. Thoracic tergum I with 3+3 chaetae, PAO with 4 vesicles, Ant. IV apical bulb simple, even anal valves each with 3 chaetae hr chilensis (Rapoport & Rubio, 1963); Chile
- 6 (5). Head with chaeta a0, Ant. IV apical bulb simple, dens each with 6 chaetae, even anal valves each with 1 chaeta hr montebella Najt & Palacios-Vargas, 1986; Mexico
- 6’. Head without chaeta a0, Ant. IV apical bulb trilobed, dens each with 5 chaetae, even anal valves each with 2 chaetae hrzerpa Weiner & Najt, 2001; Venezuela
- 7 (1’). Tibiotarsi with capitate chaetae8
- 7’. Tibiotarsi without capitate chaetae17
- 8 (7). Tibiotarsi with 7–18 capitate chaetae9
- 8’. Tibiotarsi with 1–4 capitate chaetae10
- 9 (8). Thoracic tergum I with 4+4 chaetae, tibiotarsi with about 17–18 capitate chaetae, PAO with 4 vesicles … tuberculata (Wahlgren, 1906); Falklands Is.
- 9’. Thoracic tergum I with 2+2 chaetae, tibiotarsi with 7 capitate chaetae, PAO with 5–6 vesicles ronderosi Najt, 1973; Argentina (Tierra del Fuego), Chile (Magellanes Province)
- 10 (8’). Tibiotarsi with 2–4 capitate chaetae11
- 10’. Tibiotarsi with 1 capitate chaeta15
- 11 (10). Thoracic tergum I with 3+3 chaetae, tibiotarsi I, II and III with 18, 18 and 17 chaetae respectively12
- 11’. Thoracic tergum I with 2+2 chaetae, tibiotarsi I, II and III with 19, 19 and 18 chaetae, respectively, of which 3, 4 and 4 are capitate platensis Najt & Massoud, 1974; Argentina, Australia, Tasmania
- 12 (11). Dens each with 3 chaetae, tibiotarsi I, II and III with 2, 3 and 3 or 3, 4 and 4 capitate chaetae, even anal valves each with 3 chaetae hr13
- 12’. Dens each with 4 chaetae, tibiotarsi I, II and III with 3, 4 and 4 capitate chaetae, respectively, even anal valves each with 2 chaetae hr globulosa Cassagnau & Rapoport, 1962; Argentina
- 13 (12). Tibiotarsi I, II and III with 3, 4 and 4 capitate chaetae14
- 13’. Tibiotarsi I, II and III with 2, 3 and 3 capitate chaetae, Ant. IV apical bulb slightly trilobed villalobosi Cassagnau & Rapoport, 1962; Brazil, Mexico, USA (Florida)
- 14 (13). Mucro about ½ length of dens, Ant. IV apical bulb trilobed barrerai Palacios-Vargas & Najt, 1981; Mexico
- 14’. Mucro reduced, mamelon-shaped, about ¼ length of dens, Ant. IV apical bulb simpleminimucronata Palacios-Vargas, & Najt, 1981; Mexico
- 15 (10’). Dens with 5–6 chaetae16
- 15’. Dens with 4 chaetae, PAO with 5–7 vesicles, Ant. IV apical bulb simple baconaoensis Gruia, 1983; Cuba, USA (Florida), Mexico
- 16 (15). PAO with 4–6 vesicles, Ant. IV apical bulb trilobed, dens each with 5–6 chaetaeneomexicana (Scott, 1960); USA (New Mexico, California, Louisiana, Texas, Colorado), Mexico
- 16’. PAO with 4 vesicles, Ant. IV apical bulb IV simple or bilobed, dens each with 6 chaetaececiliae Fernandes & Mendonça, 2004; Brazil
- 17 (7’). PAO with 4 vesicles18
- 17’. PAO with more than 4 vesicles25
- 18 (17). Thoracic tergum I with 3+3 chaetae19
- 18’. Thoracic tergum I with 2+2 chaetae20
- 19 (18). Ant. IV apical bulb trilobed, femora I, II and III with 13, 12 and 11 chaetae, tibiotarsi with 2 long inner chaetae B4 and B5, curved at the tipvictoriensis Izarra, 1972; Argentina
- 19’. Ant. IV apical bulb simple, femora I, II and III with 12, 11 and 10 chaetae, all tibiotarsi chaetae acuminate and straight mataraniensis Weiner & Najt, 2001; Peru
- 20 (18’). Abdominal tergum I with 1+1 sensory chaetae s, tibiotarsi I, II and III with 18, 18 and 17 chaetae21
- 20’. Abdominal tergum I with 2+2 sensory chaetae s, tibiotarsi I, II and III with 19, 19 and 18 chaetae22
- 21 (20). Head with chaeta a0, Ant. IV apical bulb trilobed, dens with 7 chaetae zapatai Najt & Palacios-Vargas 1986; Mexico
- 21’. Head without chaeta a0, Ant. IV apical bulb simple, dens with 6 chaetae stachi Mills, 1934; USA (Iowa, Connecticut, Florida Massachusetts, New York, Indiana, Missouri, North Carolina), the French West Indies, Mexico, French Guiana, Ecuador
- 22 (20’). Abdominal terga II and III with 1+1 sensory chaeta s, dens each with 5 normal chaetae23
- 22’. Abdominal terga II and III with 2+2 sensory chaetae s, dens each with 6 chaetae: 3 strong chaetae and 3 ordinary ones, even valves each with 3 chaetae hrnana Rubio & Najt, 1979; Chile
- 23 (22). Ant. IV apical bulb trilobed24
- 23’. Ant. IV apical bulb simple, even anal valves each with 2 chaetae hr, animal of a big sizeagrosa Wray, 1953; Puerto Rico, the French West Indies, Ecuador, Cuba, Brazil, French Guiana
- 24 (23). Ratio mucro : dens = 1 : 1.95, even anal valves each with 1 chaeta hr, subcoxae 2 I, II and III with 0, 1, 1 chaetae purma Weiner & Najt, 2001; Peru
- 24’. Ratio mucro : dens = 1 : 2.67, even anal valves each with 2 chaetae hr, subcoxae 2 I, II and III with 0, 2, 2 chaetae nordestina sp.n. Souza, Bellini & Weiner; Brazil
- 25 (17’). Head without chaeta a026
- 25’. Head with chaeta a027
- 26 (25). Ant. IV apical bulb trilobed, even anal valves each with 1 chaeta hr, straight mucro with apex slightly hookeddesutterae Weiner & Najt, 2001, Peru
- 26’. Ant. IV apical bulb simple, even anal valves each with 2 chaetae hr, bent mucrocontorta Denis, 1931; Costa Rica, Jamaica, Cuba, the West Indies, Hawaii, Galapagos, Mexico, India, the Cape Verde Is., Philippines, Ivory Coast, Angola
- 27 (25’). Furca well developed, dens with 5–6 chaetae, mucro present28
- 27’. Furca reduced, dens with 3–4 chaetae, mucro absent, Ant. IV apical bulb simple, PAO with 7–10 vesicles, even anal valves each with 2 chaetae hr cyanea (Rapoport, 1962); Argentina
- 28 (27). Dens with 5 chaetae 29
- 28’. Dens with 6 chaetae, Ant. IV apical bulb simple, PAO with 5–7 vesicles, even anal valves with 3 chaetae hr gabrielae Najt & Palacios-Vargas, 1986; Mexico
- 29 (28). Ant. IV apical bulb trilobed30
- 29’. Ant. IV apical bulb simple, PAO with 6–7 vesicles, even anal valves with 2 chaetae hrgrootaerti Najt, Thibaud & Jacquemart, 1991; Galapagos
- 30 (29). Even anal valves each with 2 or 3 chaetae hr31
- 30’. Even anal valves each without chaetae hr, subcoxae 2 I, II and III with 0, 2 and 2 chaetae pefauri Weiner & Najt, 2001; Venezuela
- 31 (30). Even anal valves each with 2 chaetae hr, ratio mucro : dens = 1 : 2.6saladaensis Weiner & Najt, 2001; Argentina
- 31’. Even anal valves each with 3 chaetae hr, ratio mucro : dens = 1 : 2.12 taxcoana Palacios-Vargas & Najt, 1981; Mexico
*Brachystomella mauriesi Thibaud & Massoud, 1983, B. parvula (Schäffer, 1896) and B. surendrai Goto, 1961 are not included in the key. The first species was insufficiently described. The specimens from the Neotropical region determined as B. parvula and B. surendrai are doubtful records and should be called B. ca. parvula and B.ca. surendrai.
Entomobryidae Tömösvary, 1882
Seirinae Yosii, 1961 sensu Zhang & Deharveng, 2015
Seira Lubbock, 1869
Type species: Seira domestica (Nicolet, 1842)
Genus Diagnosis
Specimens variably pigmented, at least eye patches dark. Head (ventrally and dorsally), trunk (dorsally) and appendages with strongly ciliate apically rounded and/or pointed scales. Dorsal ciliate macrochaetae variable; Abd. I macrochaetae present or absent. Antennae with 4 segments (Ant. I and II undivided); Ant. IV simple or annulated, with apical bulb unilobed, apically rounded or constricted (“heart shaped”). Prelabral chaetae smooth or ciliate, labral smooth. PAO absent. Eyes 8+8. Th. II–Abd. V trunk sensilla formula per half tergum mostly 11/022–3; Th. II–Abd. V trunk microsensilla formula per half tergum mostly 10/10100. Mesonotum variably developed. Sexual dimorphism concerning spines on first pair of legs of males usually absent; if present, Abd. I with macrochaetae. Trochanteral organ present and well developed, with several short smooth chaetae. Anterior and posterior pretarsal chaetae present. Ungues teeth well developed; unguiculus present. Abd. II–IV bothriotricha formula per half tergum: 2/3/3, rarely 2/3/2. Abd. IV at least 2.5 times longer than Abd. III in the midline. Tenent hairs capitate. Ventral tube usually with scales. Furca present and well developed, ventrally scaled (manubrium and dens), ventral manubrium with few chaetae. Dens without spines. Modified blunt macrochaetae present or absent on manubrial plates and dorsal dens. Mucro short and falcate, lacking mucronal spine (adapted after Yosii 1961; Mari Mutt 1986b; Bellini & Zeppelini 2011; Cipola et al. 2014a, b; Godeiro & Bellini 2014, 2015; Zhang & Deharveng 2015).
Lepidocyrtinus Börner, 1903 sensu Yosii, 1959
Subgenus type species: Seira (Lepidocyrtinus) annulicornis (Börner, 1903)
Subgenus diagnosis
Abd. IV with 3+3 bothriotricha. Dorsal proximal dens (and sometimes dorsal distal manubrium) with modified blunt macrochaetae (adapted from Yosii 1959).
Seira (Lepidocyrtinus) dapeste sp.n. Santos & Bellini
Habitus of Seira (Lepidocyrtinus) dapeste sp.n. (specimen fixed on ethanol, lateral view).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. antennae and head: 24, Ant. IV apical region; 25, alternative morphology of Ant. IV apical bulb; 26, Ant. III apical sense organ (lateral view); 27, left eye patch; 28, dorsal head chaetotaxy (legends to symbols in Fig. 34).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. head: 29, labral papillae region; 30, labral and prelabral chaetotaxy; 31, labial region (left side); 32, left maxillary outer lobe and sublobal plate; 33, post-labial chaetotaxy along cephalic groove.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. trunk chaetotaxy (left side): 34, legends to symbols used in dorsal chaetotaxy schemes; 35, Th. II; 36, Th. III; 37, Abd. I; 38, Abd. II.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. trunk chaetotaxy (left side): 39, Abd. III; 40, Abd. IV; 41, Abd. V.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. abdominal chaetotaxy (left side): 42, Abd. II; 43, Abd. III.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. legs: 44, trochanter III anterior side; 45, trochanteral organ; 46, leg III; 47 empodial complex I (posterior view); 48 empodial complex III (posterior view).
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
Seira (Lepidocyrtinus) dapeste sp.n. abdominal appendages: 49, ventral tube anterior face; 50, ventral tube left lateral flap; 51, ventral tube posterior face, distal region; 52, tenaculum; 53, manubrial plate chaetotaxy; 54, alternative manubrial plate chaetotaxy (left side); 55, manubrium ventral face, distal chaetotaxy; 56, dorso-distal manubrium and dorso-proximal dens modified blunt macrochaetae; 57, distal dens and mucro.
Citation: Insect Systematics & Evolution 50, 3 (2019) ; 10.1163/1876312X-00002192
ZooBank: http://zoobank.orge8f11d4a-65c4-478d-b581-3bb0ad432b19/
Type material
Holotype female, Brazil, Rio Grande do Norte State, Natal Municipality, Mata do Sagui, Campus Central da Universidade Federal do Rio Grande do Norte (5°50’34.30”S 35°12’04.63”W), 08-ix-2015, N. Santos, P. Souza & M. Freire coll., on slide without number, deposited in CC/UFRN. Paratypes: 2 males and 12 females, same data as holotype; 1 male and 1 female, same data as holotype, except collection date (03-vi-2015), all on slides without number deposited in CC/UFRN. Other paratypes: 3 females, same data as holotype, deposited in ISEA.
Diagnosis
Head partially pigmented, Ant. I-III with dark and yellow bands of pigment and leg III with two darks bands (trochanter and distal femur). Dorsal head chaetotaxy with: 7–11 An, 2 A, 3 M, 5–7 S (S1 and S3 absent in only one specimen) macrochaetae. Labral chaetae p0–1 longer than others. Ventral cephalic groove with 3+3 marginal ciliate chaetae, last pair longer than others. Th. II with 14–19 a (excluding the anterior collar), 7 m and 12–13 p macrochaetae. Th. III with 3 a, 1–2 m and 5–6 p macrochaetae. Abd. I generally with 4 (m2i–4i), Abd. II with 4 (a2, m3-3e), Abd. III with 1 (m3) central macrochaetae respectively. Abd. IV with 13–16 central (4–6 A, 2 Ae, 4–5 B and 3 C) and 11–12 lateral (5 E, 1 Ee, 4 F and 1–2 Fe) macrochaetae, with 8+8 microchaetae on posterior row. Trochanter III anterior side with 5 small spines; trochanteral organ with 24 smooth long chaetae. Tibiotarsus III external side with 4–5 long ciliate chaetae. Ventral tube anterior face with 2+2 large distal chaetae and about 7+7 strong proximal spines; posterior face with 2+2 distal smooth chaetae plus at least 5+5 small spine-like chaetae. Manubrial plate with 2+2 to 3+3 blunt macrochaetae. Dens with 4–8 blunt macrochaetae.
Description
Body pale yellow to white; proximal and distal parts of Ant. I to III, latero-posterior head, distal trochantera and femora and latero-posterior Abd. IV dark blue; eye patches dark with or without posterior pigment line; medium antenomeres, lateral head and tibiotarsi dark yellow (Fig. 23). Apically rounded strongly ciliate scales present on Ant. IV proximal part, Ant. III–I, dorsal and ventral head, dorsal trunk, all legs segment, anterior and posterior ventral tube and furca (dorsal and ventral manubrium and ventral dens). Thorax bent between Th. II and Th. III; Mesonotum elongate, projecting over posterior head (Fig. 23). Body (head + trunk) length of holotype 3390 μm, paratypes maximum body length, 3460 μm, minimum 1580 μm, average 2470 μm.
Head. Antennae smaller than the body length (holotype antennae with 2600 μm), holotype antennal ration as I: II: III: IV = 1: 1.16: 1.14: 1.98. Ant. IV weakly annulated on apex, with at least two types of chaetae: ciliate and blunt sensilla of different sizes (Fig. 24); apical bulb simple, apically constricted (Fig. 24) or rounded (Fig. 25). Ant. III apical sense organ with 2 small sensory rods in cavity, 3 surrounding guard sensilla, plus some blunt sensilla and ciliate chaetae of different sizes (Fig. 26). Eyes 8+8, largest lenses ‘A’ and ‘B’, smallest ‘G’ and ‘H’; 5 interocular chaetae present, p as macrochaeta (Fig. 27). Dorsal head chaetotaxy as in Fig. 28 (legends to symbols in Fig. 34) with: 7–11 An, 2 A, 3 M, 5–7 S (S1 and S3 absent in only one specimen) and 1 Pa macrochaetae per half head capsule; Ps5 as mesochaeta (present or absent); post-ocular bothriotrichum (Pa6) present. Two labral papillae with a spine like projection on each (Fig. 29); Four prelabral ciliate chaetae (pl1–2) (Fig. 30); labral chaetotaxy formula with 5 (p0–2), 5 (m0–2), 4 (a1–2) smooth chaetae, p0–1 longer than others (Fig. 30). Labial palp with 5 main papillae (A–E) with 0, 5, 0, 4, 4 guard chaetae, respectively; papilla E with lateral process (l.p.) thin and pointed, not reaching the papilla base; papilla H with 2 accessorial chaetae; labial papillae with 5 smooth proximal chaetae (Fig. 31). Maxillary outer lobe with 1 apical appendage and 1 subapical chaeta, both smooth; sublobal plate with 3 chaetae-like appendages (Figs. 31–32). Labial basolateral and basomedial fields with M1–2, E, L1–2 as ciliate chaetae, A1–5 and r as smooth chaetae; r chaeta reduced (Fig. 31). Post-labial region with scales and chaetae, ventral groove with 3+3 marginal ciliate chaetae, last pair longer than others (Fig. 33).
Trunk dorsal chaetotaxy (given to half tergum), legends to dorsal head and trunk chaetotaxy in Fig. 34. Th. II–Abd. V dorsal sensilla and microsensilla formulae 11/011–3 (as not seen in Abd. II and III) and 10/10100, respectively. Th. II with 14–19 a (excluding the anterior collar), 7 m and 12–13 p macrochaetae; al sensillum and ms present on lateral edge (Fig. 35). Th. III with 3 a, 1–2 m and 5–6 p macrochaetae; m6p2 as mesochaeta; al sensillum present on lateral edge (Fig. 36). Abd. I with 4 central macrochaetae (m2i–4i) – abnormally one specimen with 4+5 (a1 present on one side) and another with 3+3 central macrochaetae (m4i as microchaeta); a1a microchaeta in different positions among different specimens; ms present (Fig. 37). Abd. II with 4 central (a2, 3 m) and 1 centro-lateral (m5) macrochaetae; p6 (present or absent) and p7 as mesochaetae; m2 and a5 bothriotricha with 3–4 and 4 surrounding fan shaped chaetae, respectively; m7 and el microchaetae in different positions among different specimens (Figs. 38 and 42); acc.p6 sensillum present, as not seen (possibly absent). Abd. III with 1 central (m3) and 3 lateral (am6, pm6 and p6) macrochaetae; 7–10 lateral mesochaetae, most of them with unclear homologies; m2, a5 and m5 bothriotricha with 3, 6–7 and 4 surrounding fan shaped chaetae respectively; al sensillum and ms present, as not seen (possibly absent) (Figs. 39 and 43). Abd. IV with 13–16 central (4–6 A, 2 Ae, 4–5 B and 3 C) and 11–12 lateral (5 E, 1 Ee, 4 F and 1–2 Fe) macrochaetae; 7–9 latero-posterior mesochaetae, two of them with unclear homologies; posterior row with 8 microchaetae; ps and as plus at least 1–3 central-posterior sensilla (possibly there are more) present (Fig. 40). Abd. V with 11 (1 a, 4 m and 6 p) macro and 6 (with unclear homologies) mesochaetae; as, acc.p4 and acc.p5 sensilla present (Fig. 41).
Legs. Trochanter III anterior side with 5 small spines (Fig. 44); trochanteral organ with 24 smooth long chaetae (not spine-like as typical of Entomobryoidea) (Fig. 45). Tibiotarsus III external side with 4–5 long ciliate chaetae, distally with one straight smooth chaeta near unguiculus (Figs. 46, 48), holotype abnormally with two smooth distal chaetae in one leg. Tenent hairs discretely rough (possibly weakly ciliate), slightly longer than ungues, anterior and posterior pretarsal chaetae present (Figs. 47–48). Ungues with 1 large dorsal, 2 large paired outer and 4 inner teeth, 2 proximal paired, 1 median and 1 distal (Figs. 47–48). Unguiculi lanceolate with outer lamella slightly serrated in legs I–II (Fig. 47) and smooth in leg III (Fig. 48).
Abdominal appendages. Ventral tube anterior and posterior faces scaled, with 2+2 large plus 3+3 slender ciliate chaetae on distal region, proximal region with 3+3 ciliate chaetae plus about 7+7 strong spines (Fig. 49); lateral flaps with about 24 ciliate and 4 smooth chaetae (Fig. 50); posterior face with 2+2 distal smooth chaetae plus at least 2+2 small spine-like chaetae (Fig. 51). Tenaculum corpus with a single discretely ciliate chaeta, rami with 4+4 teeth (Fig. 52). Manubrial plate (dorsal) with 3+3 pseudopores, 2+2 to 3+3 blunt macrochaetae (2+2 in holotype) and 4+4 to 9+9 normal chaetae (Figs. 53–54 and 56); manubrium ventral face with 1+1 subapical slender (sensu Christiansen & Bellinger 2000) plus 7+7 apical normal chaetae (Fig. 55). Dens with 4–8 blunt macrochaetae, holotype with 4+4 (Fig. 56); dorsal face typically crenulate; dens lacking spines (Fig. 57). Mucro short and falcate, lacking mucronal spine (Fig. 57).
Etymology
“da peste” is a regional expression in Brazilian Portuguese, which means a brave, fearless person. It is commonly used in the northeastern region of the country.
Habitat
See Habitat section of Brachystomella nordestina sp.n. description. Specimens of Seira (Lepidocyrtinus) dapeste sp.n. were also collected during dry and rainy seasons.
Remarks
The new species resembles S. nigrans (Arlé, 1959), S. prodiga (Arlé, 1959), S. xinguensis (Arlé, 1959), S. harena Godeiro & Bellini, 2014 and S. diamantinae Godeiro & Bellini, 2015 in terms of long antennae (almost the size of trunk or longer), large and somewhat bent thorax, with mesonotum projected over head and the presence of modified blunt macrochaetae on dorsal furca. Seira (Lepidocyrtinus) dapeste sp.n. differs from S. nigrans in color pattern (the late present head totally pigmented); and from S. harena, S. diamatinae and S. prodiga specially in dorsal head microchaeta A5 (macrochaeta in the three species), macrochaeta S0 (microchaeta in S.harena) and mesochaeta Ps5 (microchaeta in S. harena, macrochaeta in S. diamantinae and S. prodiga), 14–19 a macrochaetae (excluding anterior collar) (7–8 in S. harena, 12 in S. diamantinae, 19–25 in S. prodiga) on Th. II, 7–9 central macrochaetae (3 in S. harena, 6 in S. diamantinae and S. prodiga) on Th. III, 4 central macrochaetae (2 in S. harena) on Abd. I, and 4 central macrochaetae (3 in S. harena) on Abd. II.
Discussion
The concept of chaetae homology used in the description of Brachystomella nordestina sp.n. combines the position of chaetae, as proposed by Yosii (1960), and chaetae shape plus relative position of surrounding elements, as proposed by Szeptycki (1979). In other words, we believe the dorsal long sensorial chaetae are homologous among species (particularly among the taxa which share the same formula) as well as most chaetae, and the morphological singularities are in absence, presence or relative position of part of chaetae in different species. For example, comparing B. contorta Denis, 1931 and B. mataraniensis Weiner & Najt, 2001, the first species presents 2+2 chaetae in ‘p’ position between both long sensilla on Abd. IV (see Weiner & Najt 2001, page 391, Fig. 7), while B. mataraniensis (as well as Brachystomella nordestina sp.n.) presents 3+3 chaetae between them on Abd. IV (see Weiner & Najt 2001, page 397, Fig. 33; and Fig. 12 of this paper). Following Szeptycki (1979), it is more plausible a pair of medial chaetae is missing in B. contorta than a more complicate scenario: another pair is missing in lateral position plus the long sensorial chaetae alternated their morphology with any other surrounding ordinary chaetae. In this sense we used the description of dorsal chaetotaxy of B. parvula (Schäffer, 1896) presented in Jordana et al. (1997) and Fjellberg (1998), with a few modifications, as a model of dorsal chaetotaxy within the genus, specially because it corresponds to detailed descriptions of the type species of Brachystomella. The study of detailed dorsal chaetotaxy of Brachystomellidae (as well as other Poduromorpha) holds a potential to assist separating the species with a similar morphology. To Brachystomella spp. the presence and the number of blunt chaetae on the ventral side of Ant. IV and modified eugenital chaetae surrounding male’s genital opening can also improve species identification and description, since both characters are stable in Brachystomella nordestina sp. n. type material.
The Neotropical species Brachystomella aspera (Börner, 1906) lacks a proper type locality. Börner’s description was based on the specimens harvested from orchids grown in Germany, in Hamburg Museum (University of Hamburg), originally collected from “São Francisco, Brazil”. “São Francisco” can be the largest river in northeastern Brazil or a municipality name (there are at least five Brazilian municipalities with the same name in far apart states). In the absence of the type material (destroyed or missing according curators of University of Hamburg, Germany), and a consistent type locality, the identity of B. aspera should be further investigated.
With the description of Brachystomella nordestina sp.n. there are now 10 species of Brachystomella recorded from Brazil, including: B. agrosa, B. aspera, B. ceciliae Fernandes & Mendonça, 2004, B. contorta Denis, B. garayae Queiroz & Weiner, 2011, B. ca. parvula, B. platensis Najt & Massoud, 1974, B. septemoculata Denis, 1931 and B. villalobosi Cassagnau & Rapoport, 1962 (Abrantes et al. 2010, 2012).
The first records of Brazilian Lepidocyrtinus were provided by Handschin (L. pulcher Handschin, 1924) and posteriorly by Arlé (1959). The last author described the genus Ctenocyrtinus Arlé, 1959 to include three species: Seira prodiga, S. xinguensis (originally described as S. prodiga xinguensis) and S. nigrans. Latter, Christiansen & Bellinger (2000) synonymized the genus with Seira, understating Yosii’s (1959) notes on Lepidocyrtinus and its particular morphology. The authors also made a new combination to L. pulcher, including the species in Drepanosira Bonet, 1942, even though L. pulcher has enlarged Th. II, scales on dens and lacks mucronal spine (Handschin 1924, Nikolas Cipola, personal communication). Since Lepidocyrtinus is easily recognized from Seira s. str. (see diagnosis of the subgenus and remarks of Seira (Lepidocyrtinus) dapeste sp.n.) and at least Neotropical species share several similarities, including dorsal head and trunk chaetotaxy (such as posterior head only with Pa5 macrochaeta and several multiplets of anterior macrochaetae on Th. II), we believe that Lepidocyrtinus is a well established group within Seira, and Yosii’s subgenera should be used in future descriptions.
With the description of Seira (Lepidocyrtinus) dapeste sp.n. there are now at least seven species which fit the subgenus recorded to Neotropical Region, all of them from Brazil.
We thank Dr. Gabriel Costa Queiroz for important considerations on identification of Brachystomella nordestina sp.n. Nerivânia Nunes Godeiro on identification of Seira (Lepidocyrtinus) dapeste sp.n. Nikolas Gioia Cipola for providing data of S. prodiga. The first author was supported by CNPq (Universal, Process # 441451/2014-4; PQ2015, Process # 301498/2015-6). The second and third authors were supported by CNPq/PIBIC/ICs scholarships.
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Corresponding author, e-mail: entobellini@gmail.com
