Tyrannoseira Bellini & Zeppelini, 2011,

Cipola, Nikolas Gioia, Morais, José Wellington De, Godeiro, Nerivania Nunes & Bellini, Bruno Cavalcante, 2019, Taxonomic revision of Brazilian genus Tyrannoseira Bellini & Zeppelini, 2011 (Collembola, Entomobryidae, Seirinae), Zootaxa 4586 (2), pp. 201-248: 203-242

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Tyrannoseira Bellini & Zeppelini, 2011


Genus Tyrannoseira Bellini & Zeppelini, 2011 

Type species. Seira raptora Zeppelini & Bellini, 2006 

Diagnosis of genus. Head dorsal macrochaetotaxy lacking M2, M 4i, S1, S4, S5, Ps2 and Pa4 mac ( Figs 3A, EView FIGURE 3, 4AView FIGURE 4, 8CView FIGURE 8). Four labral papillae, inner papillae bifurcate distally ( Figs 4BView FIGURE 4, 8EView FIGURE 8). Basomedian labial field with chaetae r reduced ( Figs 4A and CView FIGURE 4, 9CView FIGURE 9). Th II with 3–4 medio-median mac (m1, m 1i, m2, m 2i) and 2–3 medio-lateral mac (m4, m 4i, m4p); groups PmA –PmC with 4 –5, 2–3 and 3–5 mac, respectively ( Figs 10AView FIGURE 10, 15AView FIGURE 15, 30AView FIGURE 30). Th III with at least 7 central mac (a2, a4–5, p1–3), a6 mac or mic ( Figs 10BView FIGURE 10, 15BView FIGURE 15, 20BView FIGURE 20). Abd I only with mic ( Fig. 10CView FIGURE 10). Abd II with 3 central mac (a2, m3, m3e), m3ep absent ( Fig. 10DView FIGURE 10). Abd III with 1 central (m3) and 3 lateral mac (pm6, p6 and p6e exclusively) lateral to the bothriotricha, am6 as mic ( Fig. 10EView FIGURE 10). Abd IV with at least 8 central mac (A3, A5, B3–6, C1, T1), A3a, B1 and B2 as mac or mic ( Figs 11AView FIGURE 11, 16AView FIGURE 16, 21AView FIGURE 21, 31AView FIGURE 31). Leg I of males internally with 1 finely ciliate trochanteral spine, femur strongly enlarged and with one subdistal group of spines heavily ciliate, tibiotarsus I armed with an inner row of finely ciliate proximal spines ( Figs 5View FIGURE 5 A–C, 12View FIGURE 12 E–F). Collophore anteriorly with proximal smooth spine-like chaetae, posteriorly with distal spines ( Figs 6BView FIGURE 6, 13AView FIGURE 13).

Remarks. The distinguishing characters of Tyrannoseira  species are the fore legs of adult males, with enlarged femora each carrying one subdistal group of inner spines. Both characters are unique for the genus while others presented in the diagnosis are present on single species or some are found in combination in species of Seira  s. lat. Although some species of the domestica  -group can also have superficially similar dimorphic legs on males (but with femoral spines only in one longitudinal row), their dorsal macrochaetotaxy is clearly different, suggesting they belong to different lineages of Seirinae ( Bellini & Zeppelini 2011  , Cipola et al. 2018b).

Morphological characters of Tyrannoseira  . The following characters are shared by all five Tyrannoseira  species and are not repeated in the descriptions.

Dorsal chaetae and structure with bothriotricha ( Figs 3A, EView FIGURE 3, 7AView FIGURE 7, 8CView FIGURE 8, 10View FIGURE 10 D–E, 11AView FIGURE 11) as densely ciliate but elongated cilia, short on head (1 subantennal and 1 post-ocellar) and elongated on abdomen; Abd II–IV formula with 2 (a5, m2), 3 (a5, m2, m5) and 3 (T2, T4, D3) bothriotricha. Bothriotrichal accessory chaeta ( Figs 2IView FIGURE 2, 7AView FIGURE 7, 10View FIGURE 10 D–E, 11View FIGURE 11 A), heavily ciliate on distal two thirds or more, with fringed aspect and heavily ciliated on Abd II–IV. Mac ( Figs 2BView FIGURE 2, 3AView FIGURE 3, 7AView FIGURE 7). Finely ciliate and apically truncate-shaped mac present on dorsal head and Th II to Abd V. Mes ( Figs 1View FIGURE 1 A–C, 7AView FIGURE 7) heavily ciliate and apically acuminate, present on body, generally in Abd IV. Mic ( Figs 2View FIGURE 2 D– J, 7AView FIGURE 7, 8CView FIGURE 8, 10–11View FIGURE 10View FIGURE 11) smooth, apically acuminate and of similar lengths on different regions of the body, but with different forms, simple, with 1 median ciliation, with 4 unilateral ciliations, or with 7 or more bilateral ciliations, present on head and AMP series of Th II to Abd V. Ordinary sens ( Figs 2CView FIGURE 2, J–K, 7AView FIGURE 7, 10–11View FIGURE 10View FIGURE 11), smooth and apically rounded of similar lengths, present on Th II–III (al), Abd II–III (as, acc.p6), Abd IV (as, ps) and Abd V (as, acc.p4, acc.p5). Th II–Abd V sens formula 1, 1 | 0, 2, 2, +, 3. Psp ( Figs 2LView FIGURE 2, 7AView FIGURE 7, 10–11View FIGURE 10View FIGURE 11), as small, circular, smooth concavity with a narrow opening, present centrally on Th II to Abd IV and on coxae and manubrial plates. Specialised microchaetae – ms ( Figs 2C, HView FIGURE 2, 7AView FIGURE 7, 10–11View FIGURE 10View FIGURE 11) as small, smooth and distally striate chaetae with rounded apex present on Th II, Abd I and III. Th II–Abd V, ms formula 1, 0| 1, 0, 1, 0, 0.

Scales ( Figs 2AView FIGURE 2, 3AView FIGURE 3, 4AView FIGURE 4, 5View FIGURE 5 A–B, 6View FIGURE 6 F–G, 7View FIGURE 7 A) heavily ciliate with short interrupted cilia, generally oval or elongated and apically rounded (rarely truncate, pointed or irregular). The cilia are toothpick-shaped and abruptly acuminate at apex. Scales present on Ant I–III, both sides of head, dorsal and lateral trunk, all legs (except pretarsus), anterior collophore and furcula ventrally to lateral margin.

Antennal segments with at least 8 types of chaetae, as shown in Fig. 7B; a –fView FIGURE 7 as sens, g –h as ciliate chaetae: type a pair of rods or clubs apical organ present laterally on Ant III; type b s mooth, elongated and slightly apically acuminate, present on Ant II–IV ( Fig. 3DView FIGURE 3;, type c s mooth, somewhat short and finger-shaped, present densely on Ant I–IV ( Figs 3View FIGURE 3 C–D); type d s mooth, short and finger-shaped, present densely on Ant I–IV ( Figs 3View FIGURE 3 C–D); type e three guard sens, smooth and spiny-like surrounding apical organ of Ant III ( Fig. 3DView FIGURE 3); type f. striated and rounded, subapical modified sens of Ant III ( Fig. 3DView FIGURE 3).; type g weakly ciliate, present on Ant I–IV ( Fig. 3DView FIGURE 3); type h heavily ciliate, present on Ant I–IV ( Fig. 3DView FIGURE 3).

Antennae shorter than body length; Ant IV longer or subequal to Ant II–III, Ant I shorter ( Fig. 1View FIGURE 1); Ant IV not annulated but slightly ringed; distal, bilobed apical bulb present ( Figs 3View FIGURE 3 B–C, 8View FIGURE 8 A). Eyes oval, 8+8 per side and with 5 interocular chaetae (q, s, p, r, t) ( Figs 3AView FIGURE 3, 8CView FIGURE 8); dorsal macrochaetotaxy ( Fig. 8CView FIGURE 8) with 6–9 ‘An’ (An1a –3), 4 ‘A’ (A0, A2–3, A5), 2 ‘M’ (M1, M4), 4 ‘S’ (S0, S2–3, S6), 4 ‘Pa’ (Pa1–3, Pa5), 2 ‘Pm’ (Pm1, Pm3), 4 ‘Pp’ (Pp1– 3, Pp5) and at least 1 ‘Pe’ (Pe3); microchaetotaxy with 1 ‘An’ (An2a), 1–2 ‘A’ (A1, A4), 3 ‘M’ (M2– 4i), 1‘S’ (S5), 2 ‘Ps’ (Ps2– 3), 1 ‘Pa’ (Pa4), 2 ‘Pp’ (Pp4, Pp6) and 1–2 ‘Pe’ (Pe5–6); chaetae S1 and S4 absent. Four prelabral ciliate chaetae of subequal length; labral formula with 4 (a1–2), 5 (m0–2), 5 (p0–2) smooth chaetae, a1 apically rounded, p0–1 larger, others subequal ( Figs 4AView FIGURE 4, 8DView FIGURE 8). Labral papillae conical and apically rounded, inner papillae bifurcate distally, outer papillae simple ( Figs 4View FIGURE 4 A–B, 8View FIGURE 8 E, 14View FIGURE 14 D). Labial palp with five main papillae (A–E) plus one hypostomal papilla (H) with 0, 5, 0, 4, 4, 2 guard appendages respectively; lateral process (l.p.) apically acuminate and surpassing the base of apical appendage ( Figs 4DView FIGURE 4, 9BView FIGURE 9). Labium with five smooth proximal chaetae (p.c.) ( Figs 4DView FIGURE 4, 9BView FIGURE 9). Basolateral and basomedian labial fields with chaetae a1–5 smooth, M1–2, E, L1–2 ciliate, r spine-like, reduced and smooth ( Figs 4A and CView FIGURE 4, 9CView FIGURE 9). Maxillary palp with apical appendage (a.a.) and basal chaeta (b.c.) slightly ciliate and subequal in size; sublobal plate internally with 3 main, smooth appendages, externally with 1 minute, smooth appendage ( Figs 4AView FIGURE 4, 9AView FIGURE 9).

Thoracic chaetotaxy ( Figs 10View FIGURE 10 A–B, 15View FIGURE 15 A–B, 20View FIGURE 20 A–B, 25View FIGURE 25 A–B, 30View FIGURE 30 A–B). Th II with 21–28 main mac (a 4i, a4+, a5ip –5p, m1– 1i, m2– 2i, m 4i –4p, p1ip2–1p, p2a –2p, p2ea2–2ep) and 10 main mic (a2p, a5ip, m 1i 2, m4ip, m5a – 5p, p4, p6–6e), m 1i present or absent, m4p rarely as mic, p1ip2 rarely as mac, p 1i 2, p1ip, p2p, p2ea and p2ea2 (present or absent) as mac or mic. Th III with 8–11 mac (a2, a4–6, m6–6p, m7, p1, p2–2a, p3) and 8–10 mic (a1a – 1, a3, a6–7, m4–5, p 1i, p4–6), a6 as mac or mic, a3 rarely as mac and p 1i present or absent.

Abdominal chaetotaxy ( Figs 10View FIGURE 10 C–E, 11View FIGURE 11 A–B, 15View FIGURE 15 C–E, 16View FIGURE 16 A–B, 20View FIGURE 20 C–E, 21View FIGURE 21 A–B, 25View FIGURE 25 C–E, 26View FIGURE 26 A–B, 30View FIGURE 30 C–E, 31View FIGURE 31 A– B). Abd I with 12 mic (a1–3, a5–6, m2–6, p5–6). Abd II with 4 mac (a2, m3, m3e, m5), 1 mes (p6), 8–10 mic (a2p, a3, a6–7, m3ea, m6–7, p5, p7, el), and bothriotricha a5 and m2 with 5–7 and 3–4 accessory chaetae, respectively; chaeta a2p rarely present, p7 and el rarely as mes. Abd III with 5 mac (m3, pm6, p6, p6e, p7), 6–8 mic (a3, a7, a8, am6, m7, m8, p5, p 7i), bothriotrichum m2 with 4–5 accessory chaetae (a1–2 and 2 unnamed, p3 accessory or mic) and bothriotricha a5 and m5 with 10–11 accessory chaetae between them (em, emp and 8 unnamed, c3 accessory or mic), a8 and m8 present or absent, p 7i mes or mic rarely absent, p8 rarely present, p3 as accessory chaeta or mic, present or absent. Abd IV with 21–29 mac (A3a –3, A5, B1–6, C1, T1, T7, D3p, E2–4p, Ee10, F1–3p, Fe2–6), about 15 mic (A1–2, A4, A6, Ae7, C1p –4, T3, T5–6, D2a –2, E1), bothriotricha T2 and T4 with 4–5 (s, m, T1p, D1–1p) and 3 (T4a, Pi, Pe) accessory chaetae, respectively, A3a, B1, B2, T6, Ee10 and F1p as mac or mic, D3p and F3p as mes or mic, C1p rarely present, m present or absent; posteriorly with 4–5 mes. Abd V with 9–14 mac (a5, m2, m3, m5, m5e, p1, p3–6e, p3pi, p5a, p6ai) and 10–16 mes or mic (a1, a3, a6, m5a, m5ea, p3a –6ae, p1p –5pe, pp6), a5 as mac, mes or mic, p5a, p6ai and p3pe as mac or mes, p4a, p5ae, p5pi, p5pe, p6e and pp6 present or absent.

Legs with subcoxa I having one row of ciliate chaetae and 2 psp; subcoxa II with two rows (anterior and posterior) of ciliate chaetae and 3 psp; subcoxa III with one row of ciliate chaetae and 2 (rarely 3) psp ( Figs 12View FIGURE 12 A– C). Leg I coxa in males with 1 subdistal, finely ciliate inner spine, apically pointed ( Fig. 12AView FIGURE 12). Trochanter with 1 inner, club-shaped spine, finely ciliate ( Fig. 5AView FIGURE 5). Femur strongly enlarged on outer and inner faces and with one subdistal group of heavily ciliate inner spines, varying in thickness and apically pointed ( Figs 5View FIGURE 5 A–B). Tibiotarsus armed by a single inner row of finely ciliate, apically rounded spines (proximal shorter, median larger and apically curved), also with heavily ciliate and apically acuminate distal spine-like mac ( Figs 5A, CView FIGURE 5, 12EView FIGURE 12); outer face distally with 1 finely ciliate, apically capitate, tenent hair, inner face of tibiotarsus III with one weakly grooves smooth chaetae on distal two thirds ( Figs 5View FIGURE 5 D–E). Pretarsus with one minute anterior and one posterior smooth chaetae ( Fig. 12GView FIGURE 12). Unguis inner face with 2 paired basal teeth, 1 unpaired median tooth, and 1 unpaired apical tooth; outer face with 2 paired lateral teeth and one unpaired apically pointed basomedian lamella ( Figs 5View FIGURE 5 D–F). Unguiculus with 4 acuminate lamellae, pe lamella serrated on distal half and with a small median tooth, other lamellae smooth (ai, ae, pi) ( Figs 5E, GView FIGURE 5, 15GView FIGURE 15). Collophore ( Figs 6View FIGURE 6 A–B, 13AView FIGURE 13, 28AView FIGURE 28) with 9–10 chaetae on anterior face, proximal region with 1 psp, 3–4 reduced spine-like smooth chaetae and at least 4 larger, ciliate, apically acuminate chaetae, plus 1–2 ciliate mac distally of equal or differing lengths; posterior face with 2–5 distal chaetae, 1–3 smooth spines, 1 ciliate chaeta present or absent plus 1 smooth chaeta; lateral flap with distal smooth chaetae and proximal ciliate chaetae. Tenaculum with 4 teeth on each ramus; corpus with one weakly ciliate chaeta. Genital plate circinate in males, with 15 circumgenital smooth chaetae (1 superior unpaired) and 3+3 small eugenital smooth chaetae ( Figs 6CView FIGURE 6, 13BView FIGURE 13). Female with two pairs (superior and inferior) of small smooth chaetae, without other modifications. Furcula ( Figs 1View FIGURE 1, 6DView FIGURE 6) subequal to length of abdomen. Manubrium shorter than dens and with ciliate chaetae; ventral face with ciliate chaetal formula specific to each species ( Figs 13CView FIGURE 13, 18BView FIGURE 18, 23BView FIGURE 23, 28BView FIGURE 28, 33BView FIGURE 33); dorsal face densely covered with ciliate chaetae of different lengths, manubrial plate distally with up to 5 ciliate chaetae and 3 psp. Dens dorsally crenulate with numerous ciliate chaetae, lacking other modifications such as spines or tubercles ( Fig. 6EView FIGURE 6). Mucro dorsally falcate, without basal spine ( Fig. 6EView FIGURE 6).

Remarks. Corrections to the character descriptions include the apical bulb of Ant IV which was originally described as absent in T. diabolica  and unilobed in other species, but it is bilobed in all species ( Figs 3CView FIGURE 3, 8AView FIGURE 8). From lateral view the apical bulb of Ant IV can be misinterpreted as having a single lobe (e.g. Zeppelini & Lima 2012: 38, fig. 2), because the lobes can overlap, and so they only the dorsal or ventral face be viewed. Other character typical of all Tyrannoseira  spp. the chaeta r is spine-like and reduced in basomedian labial field, but it was reported as absent in T. diabolica  , ciliate in T. gladiata  and smooth in T. raptora  . It is clear in the MEV photograph that the r chaeta is reduced and partially obscured by the a2 smooth chaeta. In the original description r chaeta was mistakenly identified as a2 chaeta (see Zeppelini & Bellini 2006: 25, right face of fig.4). Similarly T. diabolica  was reported to have 3 inner teeth on the ungues, but it bears 4 teeth as do all other species.

A detailed chaetotaxy for all species, as done here, needed to be described so that they could be confidently compared. On the head there are 4 mac on ‘A’ series (A0, A2, A3, A5), but A5 was originally omitted in the descriptions of T. raptora  and T. sex  (see Zeppelini & Bellini 2006: 27, fig. 8a; Bellini & Zeppelini 2011: 553, fig. 6), and two A0 was a misinterpretation of the frontal head of T. gladiata  (see Zeppelini & Lima 2012: 41, fig. 17)., The postocular mac (Pa5 and Pp5), as well as the five posterior mac (Pa1, Pm1, Pp1, Pp2, Pe3) also dorsally on the head, were partly omitted from the descriptions, except for T. gladiata  . On Th II anteriorly, T. raptora  may have 1 extra mac (a5+), while T. diabolica  , T. gladiata  and T. sex  have 2 extra mac (a4+ and a 4i). On Abd II there are 3 central mac (a2, m3 and m3e) in all species ( Figs 10DView FIGURE 10, 15DView FIGURE 15, 20DView FIGURE 20, 25DView FIGURE 25, 30DView FIGURE 30), but T. bicolorcornuta  was previously reported with 5 and T. gladiata  with 2 mac (see Zeppelini & Lima 2012: 41, fig. 17). Laterally on Abd III there are 4 mac typically present (pm6, p6, p6e and p7), while only 2 or 3 lateral mac have been reported for all species except for T. bicolorcornuta  ) in this region. As for Abd IV macrochaetotaxy, T. bicolorcornuta  and T. diabolica  were reported to have 12 and 13 central mac, respectively. However, number of central mac on Abd IV is at most 10 including 8 central mac (A3, A5, B3–6, C1, T1) which is constant for the genus (e.g. Fig. 11AView FIGURE 11), and further mac (A3a, B1–2, T7) are constant for specific species only ( Figs 16AView FIGURE 16, 21AView FIGURE 21, 26AView FIGURE 26, 31AView FIGURE 31). The B2 mac is constant only in T. bicolorcornuta  , T. gladiata  and T. sex  , while the other mac can have intraspecific variations that are present or absent as A3a in T. diabolica  , B 1 in T. sex  , and T 7 in T. gladiata  . Other variations thatwere not reported in the original descriptions follow. In ‘m’ series of Th II, m 4i mac was observed as present or absent in T. raptora  , while m 1i in T. bicolorcornuta  can be present or absent as well. On the same segment, in the PmA –PmC group, the mac that may be present or absent are: p1ip 2 in T. sex  , p2ea in T. raptora  , p 1i 2 and p2p in T. bicolorcornuta  , and p2ea 2 in T. gladiata  and T. sex  ( Figs 10AView FIGURE 10, 15AView FIGURE 15, 20AView FIGURE 20, 25AView FIGURE 25, 30AView FIGURE 30). Note that out of these variations on both segments (Th II and Abd IV), only the latter (p2ea 2 in Th II) is variable in more than one species, and for this reason the remaining variations can help in the species diagnoses.

Other characteristics omitted from the original descriptions that are now useful to separate species are: Ant III modified subapical sens, labral papillae shape, chaetotaxy of clypeus, ventral head, collophore, manubrial ventral formula and manubrial plate (see Table 1).

Distribution and habitat of Tyrannoseira  . The genus has so far only been found in the northeastern region of Brazil, from Ceará (new record), Paraíba, Pernambuco and Rio Grande do Norte states ( Fig. 34View FIGURE 34), i.e. Good’s biogeographic zone 27 of the Neotropical region ( Good 1974). The regional climate is tropical with a dry season (Aw) and is generally hot semi-arid (BSh), as in northern Atlantic Rainforest (low humidity) and Caatinga (low rainfall) biomes, respectively ( Kottek et al. 2006).

Tyrannoseira  is apparently endemic to this region of Brazil, but its preferred microhabitat is still uncertain.

Most species were found living among granitic rocks, even at times of high solar incidence, i.e. above 40°C in the Caatinga phytogeographic domain. Only T. bicolorcornuta  (in part) and T. diabolica  were found in Atlantic Rainforest. The northern Atlantic Rainforest of Neotropical region suffers from the strong influence of Caatinga, which may be the reason these latter two species occur there.