Research Article |
Corresponding author: Bruno Cavalcante Bellini ( entobellini@gmail.com ) Academic editor: Louis Deharveng
© 2023 Bruno Cavalcante Bellini, Mariana Fernandes De Oliveira, Wanda Maria Weiner, Rudy Camilo Nunes, Gleyce Da Silva Medeiros.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bellini BC, Oliveira MFD, Weiner WM, Nunes RC, Medeiros GDS (2023) Revisiting Szeptyckitheca Betsch & Weiner (Collembola, Symphypleona, Sminthuridae): new species, updated diagnoses, and a key. ZooKeys 1186: 139-174. https://doi.org/10.3897/zookeys.1186.111837
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Szeptyckitheca Betsch & Weiner is a genus of Sphyrothecinae (Sminthuridae) similar to Sphyrotheca Börner, with 13 nominal species. Most descriptions of Szeptyckitheca taxa lack valuable data in face of the current taxonomy of Symphypleona. In this study the previously described species of the genus were surveyed, aiming to provide updated diagnoses for them. Three species were also transferred to Szeptyckitheca: Sphyrotheca karlarum Palacios-Vargas, Vázquez & Cuéllar, 2003, S. peteri Palacios-Vargas, Vázquez & Cuéllar, 2003, and S. koreana Betsch & Weiner, 2009, based on trochanteral and/or dental chaetotaxy. Two new Brazilian species of the genus are described and illustrated, S. andrzeji Medeiros, Bellini & Weiner, sp. nov., with a remarkable reduced ventral dental chaetotaxy not seen in other Neotropical species, and S. cyanea Oliveira, Medeiros & Bellini, sp. nov. with a distinctive large set of head vertex spines (18). Finally, an updated key to all the valid species of the genus is presented.
Chaetotaxy, Neotropical fauna, Sphyrotheca Börner, Sphyrothecinae, taxonomy
Szeptyckitheca Betsch & Weiner, 2009 is the second largest genus of the small subfamily Sphyrothecinae (Sminthuridae) with 13 described species, after Sphyrotheca Börner, 1906 with 21 (
Szeptyckitheca has a mainly Holotropical distribution, with few species occurring from other regions but under subtropical climates (
Here we described in detail two new species of Szeptyckitheca from Brazil. We surveyed previously described taxa of the genus to present the current state of the knowledge of the group. We also transferred Sphyrotheca karlarum Palacios-Vargas, Vázquez & Cuéllar, 2003, S. peteri Palacios-Vargas, Vázquez & Cuéllar, 2003, and S. koreana Betsch & Weiner, 2009 to Szeptyckitheca. With this study we were able to present update diagnoses of Szeptyckitheca and its species, provide some taxonomic notes, and an updated identification key for them.
The studied specimens of Szeptyckitheca were preserved in 70% ethanol at 6 °C, clarified in Nesbitt’s solution, washed in Arlé’s liquid, and mounted on glass slides in Hoyer’s medium, combining procedures outlined by
For our survey of the previously described Szeptyckitheca species, we consulted the original descriptions as well as the redescription of S. santiagoi (Yosii, 1959) by
The terminology used in the diagnoses and descriptions follows
Abbreviations used in the descriptions and figures are: Abd. —abdominal segment(s); Ant. —antennal segment(s); mac —macrochaeta(e); Th. —thoracic segment(s).
On the figures, chaetae present or absent are marked with white arrows; unpaired chaetae on head and trunk are marked with an asterisk, *. Ant. IV subsegments are counted from the base to the apex. The head vertex was considered as the frontal and interantennal areas sensu
Head, trunk (thorax + abdomen), and furca chaetotaxy are given by half body, with the exception of head vertex chaetae which are listed as their total number. Chaetae labels are marked in bold.
The type series of the new species are deposited at the Collembola Collection of the Biosciences Center of the Federal University of Rio Grande do Norte (
Order Symphypleona Börner, 1901 sensu Bretfeld, 1986
Suborder Appendiciphora Bretfeld, 1986
Superfamily Sminthuroidea Bretfeld, 1994
Family Sminthuridae Lubbock, 1862
Subfamily Sphyrothecinae
Cuticle granulation rough. Specimens pigmented, eyepatches dark. Ant. IV with 8–12 subsegments. Eyes 8+8, head without vesicles or tubercles, eyepatches with 0–2 interocular chaetae, head vertex with a total of 4–18 strong erect large spines, cephalic chaeta A1 present or absent. Large abdomen without mesothoracic vesicles, dorsally with spine-like, blunt, or curved mac, adults with bothriotricha A–D, parafurcal area with 1+1 neosminthuroid chaetae. Small abdomen without bothriotrichum E, female subanal appendage short (not reaching the apex of the ventral anal valves), long (surpassing the ventral anal valves) or very long (surpassing the dorsal anal valve). Trochanteral spines present on legs I and III, sometimes on leg II as well, ungues usually with the internal tooth, pseudonychia and tunica. Manubrium with six or seven dorsal chaetae. Ventral (anterior) dental chaetotaxy with two to four transversal rows of chaetae, with the following resumed chaetotaxy formula from the apex to the base: 4–2,2–0,2–0…1, dorsal dens with 13–24 chaetae. Mucro without chaeta, with an apical notch (data based on
Szeptyckitheca kesongensis Betsch & Weiner, 2009.
The closest genus to Szeptyckitheca is Sphyrotheca, and many features in their diagnoses overlap (
Species / features | S. andrzeji sp. nov. | S. cyanea sp. nov. | S. bellingeri | S. kac | S. karlarum | S. mucroserrata | S. peteri | S. vanderdrifti |
---|---|---|---|---|---|---|---|---|
Trunk color pattern | pinkish, with purple dorsal spots | dark bluish | dorsally dark brownish with lateral sides bluish | with dark spots | ♂ pale bluish with purple spots and stripes, ♀ yellowish with brown spots and stripes | with lateral weak purple bands, posteriorly purple | ♂ pale bluish with purple spots, ♀ yellowish with brown spots | yellowish with indistinct areas of pigment, or brownish |
Ant. IV subsegments | 11 | 11 | 8 | 9 | 9–10 | 9–10 | 9–10 | ~10 |
Ant. III chaetae | 19 | 21** | 18 | 14 | 19 | ? | 17? | 19 |
Ant. II chaetae | 13 | 15 | 14? | 13 | 15 | ? | 12 | 14 |
Ant. II long chaetae | 3 | 4 | 3 | 3 | 2 | ? | 2 | 2–3 |
Ant. I chaetae | 6 | 6 | 5? | 7 | 6 | ? | 4? | 6 |
Interocular chaetae | - | 2 | 2 | - | 2 | 2 | 1 | 2 |
Frontal head A1 | - | + | - | - | - | - | - | - |
Head vertex large spines (total) | 14 | 18 | 14 | 16 | 16 | 16 | 16 | 16 |
Head vertex rough spines | - | - | -? | -? | + | + | + | - |
Head frons sec. reduced chaetae | + | - | + | + | + | -? | + | + |
Head interantennal bifid chaetae (♀) | - | - | - | - | - | - | - | - |
Head clypeal f line mac | - | - | - | 4 | - | - | - | - |
Tr. I n° of spines | 2 | 2 | 2 | 1 | 1 | ? | 1 | 1 |
Tr. I spine(s) apex shape | capitate | capitate | acuminate | capitate | blunt | ? | ? | acuminate |
Tr. II spine apex shape | blunt | capitate | blunt | knobbed | - | ? | knobbed | acuminate? |
Tr. III spine apex shape | blunt | blunt | blunt | blunt | blunt | blunt | blunt | capitate |
Tr. III regular chaetae | 5 | 5 | 5 | 3 | 5 | 5 | 5 | 4 |
Tr. III oval organs | 1 | 1 | -? | - | 1 | + | 1 | -? |
Femur I spine | +(1) | +(1) | +(1) | +(1) | +(1) | ? | +(1) | ? |
Ungues inner tooth | + | + | + | + | + | + | + | + |
Ungual tunica | + | + | + | + | + | + | + | + |
Ungual pseudonychia | weak | weak | strong | weak | + | - | + | strong |
Unguiculus I internal tooth | - | + | - | + | + | - | + | + |
Unguiculus III apical filament | thin | thin | thick | thin | thin | thin | thin | thin |
Unguiculus III filament length compared to unguis III | = | < | > | = | < | = | > | < |
Large abdomen capitate mac | + | + | - | + | + | - | +* | - |
Female anal valve mps1 winged | - | - | - | - | - | - | - | - |
Female subanal appendage shape | short, spoon-like, apically serrated | long, slightly curved at the apex, unilaterally serrated | long, slightly curved at the apex, smooth or unilaterally serrated | long, blunt, apically serrated | long, spatulated, serrated at the tip | long, spatulated or acuminated, serrated at the tip | long, spatulated, serrated at the tip | long, acuminate, apically or almost entirely serrated |
Manubrial chaetae | 7 | 7 | 6 | ? | ? | 7 | ? | ? |
Dens ventral chaetotaxy | 3…1 | 3,2…1 | 2,2…1 | 4,1...1 | 3,2…1 | 3,2…1§ | 3,2…1 | 2,2…1 |
Dens dorsal chaetae | 17 | 16 | 17 | 15 | 12 | 14§ | 17 | 16 |
Prominent mucronal notch | - | + | + | + | - | - | - | - |
Species / features | S. boneti | S. coerulea | S. formosana | S. implicata | S. kesongensis | S. koreana | S. machadoi | S. nepalica | S. santiagoi | S. spinimucronata |
---|---|---|---|---|---|---|---|---|---|---|
Trunk color pattern | dorsally black or purplish-red | bluish with white spots | dorsally yellowish with lateral sides purplish | dark yellowish with purple stripes and spots | pale | lateral large abdomen violet | with transversal stripes and spots of dark pigment | pale, with diffused pigment and violet patches | variable, usually laterally dark and dorsally pale | whitish, with diffused fields of purple pigment |
Ant. IV subsegments | 10–12 | 9–10 | 9–10 | 11 | 10 | 9 | 10 | 10 | 10 | ~10 |
Ant. III chaetae | 24* | ? | ? | ? | 23 | 23 | 21* | ? | 15? | ? |
Ant. II chaetae | ? | 15–16 | ? | >8 | 16 | 16 | 15* | ? | 15? | ? |
Ant. II long chaetae | ? | -? | ? | - | - | - | 4 | -? | 1 | ? |
Ant. I chaetae | ? | 7 | 4? | 5 | 7 | 7 | 5? | ? | 5 | ? |
Interocular chaetae | 2 | 2 | -? | 1 | 2 | 2 | 2? | -? | 1 | -? |
Frontal head A1 | + | + | + | + | - | + | + | + | - | + |
Head vertex large spines (total) | 10 | 6 | 15 | 16 | 16 | 4**** | 16 | 11 | 14 or 16 | 16 |
Head vertex rough spines | -? | - | + | + | +/- | +/- | -? | -? | +? | + |
Head frons sec. reduced chaetae | + | - | -? | - | - | + | - | -? | +? | - |
Head interantennal bifid chaetae (♀) | - | - | - | - | + | - | - | - | - | - |
Head clypeal f line mac | - | - | - | - | - | - | - | - | - | - |
Tr. I n° of spines | ? | 1 | ? | 1 | 1 | 1 | 1 | 1 | 1 | ? |
Tr. I spine apex shape | ? | blunt | ? | blunt | capitate | capitate | blunt | blunt | blunt | ? |
Tr. II spine apex shape | ? | blunt | ? | blunt | capitate | capitate | - | blunt | blunt | ? |
Tr. III spine apex shape | ? | blunt | ? | blunt | capitate | capitate | blunt | blunt | blunt | blunt |
Tr. III regular chaetae | ? | 5 | ? | 5 | 5 | 4 | 4 | ? | 5 | 5 |
Tr. III oval organs | ? | -? | ? | -? | 2 | 2 | - | ? | ? | -? |
Femur I spine | ? | +(1) | ? | ? | +(1) | +(1) | +(1) | ? | +(2) | ? |
Ungues inner tooth | + | + | + | + | + | + | + | + | - | + |
Ungual tunica | -** | + | + | + | + | + | + | + | + | + |
Ungual pseudonychia | weak | strong | - | weak | strong | strong | strong | strong | weak | - |
Unguiculus I internal tooth | + | - | + | + | - | + | + | +/- | +/- | - |
Unguiculus III apical filament | thin | thin | thin | thin | thin | thin | thin | thin? | thin | thin |
Unguiculus III filament compared to unguis III | < | < | < | < | < | = | > | < | = | < |
Large abdomen capitate mac | ? | + | - | + | - | - | ? | - | - | - |
Female anal valve mps1 winged | - | - | - | - | + | - | - | - | - | - |
Female subanal appendage shape | very long, acuminate, serrated at the middle | long, acuminate, unilaterally serrated | long, blunt, apically serrated | long, acuminate, apically serrated | long, acuminate, apically serrated | long, blunt, apically unilaterally serrated | long, spatulated, apically serrated | long, acuminate, apically serrated | long, with a bidentate apex, smooth or apically serrated | long, acuminate, apically serrated |
Manubrial chaetae | ? | 7 | 7 | 7 | ? | ? | ? | 6 | ? | 7 |
Dens ventral chaetotaxy | 3,2,2…1 or 3,2…1 |
4,2…1 | 3,1…1 | 2,2…1 | 3,2…1 | 3…1 or 2…1 |
2,2…1 | 3,2....1 | 2,2…1*** | 3,2…1 |
Dens dorsal chaetae | 17–18 | 16 | 15 | 19 | 20 | 17 | 24 | 13 | 13*** | 17 |
Prominent mucronal notch | + | + | + | + | + | + | + | + | + | + |
In our survey and the descriptions of the new species we observed some species hold reduced spines on the frontal head. We considered such reduction as secondary, since other taxa of the genus present all frontal head chaetae well developed, a feature also seen in other genera and families of Symphypleona (
Sphyrotheca bellingeri Betsch, 1965: 444.
Yellowish ground, head with dark blue bands dorsally in a V-shape, dorsal trunk dark brownish with bronze, bluish or yellowish effects, other regions including appendages bluish. Ant. IV with 8 subsegments; Ant. III with 18 chaetae other than the sensory clubs; Ant. II with three chaetae clearly longer than the others. Eyepatches with two interocular chaetae modified into strong spines each. Head vertex with a total of 14 large spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaeta near the spines present. Trochanters I–III with 2,1,1 spines, respectively, trochanter I spines acuminate, trochanters II and III spines blunt; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thick and surpassing the tip of the unguis III. Large abdomen lacking capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), slightly curved at the apex (hook-like), smooth or with the external border serrated. Manubrium with 6+6 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 2,2…1, dorsal chaetotaxy with 17 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca bellingeri is the sole species of the genus with eight subsegments of the Ant. IV, while all others have nine or more. It also shares with the two new described species a pair of trochanteral spines on leg 1, a feature not reported in any other taxon of the genus (see Tables
Specimens were found on mosses and liverworts growing on logs and stony ground, over the vegetation, litter layer, dead branches and directly upon the soil (
Jamaica (
Sminthurus Boneti [sic] Denis, 1948: 298.
Dorso-posterior head and dorsal trunk blackish to purplish-red, legs and antennae weakly pigmented, ventral side pale. Ant. IV with 10–12 subsegments; Ant. III with 24 chaetae other than the sensory clubs, including two peculiar small sensilla within cavities. Eyepatches with two regular interocular chaetae each. Head vertex with a total of ten large spines, two of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines present. Ungues with a single inner tooth, without tunica (unguis III possibly with a small distal rudiment of tunica) and with a weakly developed pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Female with a very long subanal appendage (surpassing the dorsal anal valve), acuminate, feathered (serrated) at the middle region on its both edges. Dens ventral chaetotaxy formula from the apex to the base as: 3,2…1 or 3,2,2…1, dorsal chaetotaxy with 17 or 18 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca boneti is unique compared to all its congeners by the very long subanal appendages of the females, surpassing the dorsal anal valves. However, we could not find any data on the species legs chaetotaxy, especially the presence and shape of the trochanteral spines. Apparently the species was revised by Betsch (
Specimens were found in bushes (
Vietnam (
Sphyrotheca coerulea Bretfeld, 2005: 32.
Head and body almost completely blue, with pale spots. Ant. IV with nine or ten subsegments; Ant. II with 15 or 16 chaetae; Ant. I with seven chaetae. Eyepatches with two regular interocular chaetae each. Head vertex with a total of six large smooth spines, none of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines absent. Trochanters I–III with 1,1,1 spines, respectively, all blunt; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen dorsally with several capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, apically serrated on its internal face. Manubrium with 7+7 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 4,2…1, dorsal chaetotaxy with 16 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca coerulea is the only species of the genus with six spines on the head vertex. Further data on the species are presented in Table
Specimens were found on shrubs and grasses near a stream of water (
Yemen (Socotra Island) (
Sphyrotheca formosana Yosii, 1965: 49.
Yellowish ground, antennae diffusely pigmented, head almost pale, large abdomen laterally purplish, dorsally pale. Ant. IV with nine or ten subsegments. Head vertex with a total of 15 large spines, three of them unpaired; unpaired chaeta A1 present. Ungues with a single inner tooth, with tunica and without pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen without capitate chaetae. Female’s subanal appendage long (surpassing the ventral anal valves), blunt, apically serrated on its external face. Dens ventral chaetotaxy formula from the apex to the base as: 3,1…1, dorsal chaetotaxy with 15 chaetae; mucronal notch prominent (adapted from
Like S. boneti, S. formosana was apparently revised by Betsch (
Unknown.
Taiwan (
Sphyrotheca implicata Hüther, 1967: 252.
Dark yellowish ground, with dark purple spots and two longitudinal stripes on the dorsal large abdomen, appendages pale purple. Ant. IV with 11 subsegments; Ant. II with more than eight chaetae, none of them clearly longer than the others; Ant. I with five chaetae. Eyepatches with one regular interocular chaeta each. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines absent. Trochanters I–III with 1,1,1 spines, respectively, all blunt; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and weak pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen dorsally with several capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, apically serrated. Manubrium with 7+7 chaetae, dens ventral chaetotaxy formula from the apex to the base as: 2,2…1, dorsal chaetotaxy with 19 chaetae; mucronal notch prominent (adapted from
Specimens were found associated to bushes (
South Sudan (
Szeptyckitheca kac Zeppelini, Lopes & Lima, 2018: 3.
Trunk with dark spots of pigment, mucro and legs less pigmented. Ant. IV with nine subsegments; Ant. III with 14 chaetae other than the sensory clubs; Ant. II with 13 chaetae, three of them clearly longer than the others; Ant. I with seven chaetae; some chaetae of Ant. II–IV capitate. Eyepatches lacking interocular chaetae. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaetae near the spines present. Clypeal area with 4+4 mac near the antennae. Trochanters I–III with 1,1,1 spines, respectively, trochanter I spine capitate, II knobbed and III blunt; trochanter III with three regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and weak pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and reaching the tip of the unguis III. Large abdomen dorsally with several capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), blunt, apically serrated on its internal face. Dens ventral chaetotaxy formula from the apex to the base as: 4,1…1, dorsal chaetotaxy with 15 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca kac is unique within the genus due the presence of 4+4 modified mac on the upper clypeus (f line) and the dens ventral chaetotaxy formula of 4,1...1 chaetae. Further data on the species are presented in Table I.
Specimens were found in the canopy of rainforest (
Brazil (
Sphyrotheca karlarum Palacios-Vargas, Vázquez & Cuéllar, 2003: 303–306, 308, figs 4–6, Mexico, Quintana Roo, Reserva de la Biosfera de Sian Ka’na (orig. descr.).
Males pale bluish, with purple or blue spots on antennae, dorsal head and furca, dorsal trunk striped; females yellowish, with brown spots and stripes with the same distribution of males. Ant. IV with nine or ten subsegments, with some proximal chaetae capitate; Ant. III with 19 chaetae other than the sensory clubs, 4–6 of them longer than the others and capitate; Ant. II with 15 chaetae, two of them clearly longer than the others, two of them modified into spines; Ant I with six chaetae. Eyepatches with two interocular chaetae each, one of them modified into a spine. Head vertex with a total of 16 large and rough spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaetae near the spines present. Trochanters I–III with 1,0,1 spines, respectively, trochanters I and III spines blunt; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen with capitate chaetae. Female with a long subanal appendage (slightly surpassing the ventral anal valves), spatulated, apically serrated on both faces. Dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 12 chaetae; mucronal notch discrete (adapted from
Sphyrotheca karlarum is herein transferred to Szeptyckitheca due to the presence of robust and mostly erect spines on the head vertex and dorsal large abdomen, presence of three transversal rows of dental ventral chaetae and presence of spines on trochanters I and III, features used by
Specimens were found in low flooded jungle (
Mexico (
Szeptyckitheca kesongensis Betsch & Weiner, 2009: 40.
Specimens pale or very clear. Ant. IV with ten subsegments; Ant. III with 23 chaetae other than the sensory clubs; Ant. II with 16 chaetae, none of them clearly longer than the others; Ant. I with seven chaetae. Eyepatches with two regular interocular chaeta each. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaetae near the spines absent. Head interantennal area with 1+1 bifid chaetae. Trochanters I–III with 1,1,1 spines, respectively, all capitate; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen without capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, apically serrated on both edges. Dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 20 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca kesongensis, the type species of the genus, shows a very peculiar head morphology, with the female bearing bifid chaetae in the interantennal area, while the male lack such morphology (
Specimens were found associated to Robinia sp., shrubs, pines, and plant debris (
North Korea (
Sphyrotheca koreana Betsch & Weiner, 2009: 36–39, figs 1–13, North Korea, Kaesong-si province, Chonma-sun Mountains (orig. descr.).
Pale background, with antennae, frontal area (interocular field) and lateral sides of large abdomen violet. Ant. IV with nine subsegments; Ant. III with 23 chaetae other than the sensory clubs; Ant. II with 16 chaetae, none of them clearly longer than the others; Ant. I with seven chaetae. Eyepatches with two regular interocular chaetae each. Head vertex with a total of four large erect spines, plus four short spines and nine rough curved mac, three of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines present. Trochanters I–III with 1,1,1 spines, respectively, all capitate; trochanter III with four regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and reaching the tip of the unguis III. Large abdomen without capitate mac, but with rough curved mac. Female with a long subanal appendage (surpassing the ventral anal valves), blunt, apically serrated on its internal edge. Dens ventral chaetotaxy formula from the apex to the base as: 3…1 or 2…1, dorsal chaetotaxy with 17 chaetae; mucronal notch prominent (adapted from
Sphyrotheca koreana is herein transferred to Szeptyckitheca due to the presence of one spine on trochanters I–III. This feature, which was listed as diagnostic of the later genus by
Specimens were found in forested areas (
North Korea (
Sminthurotheca machadoi Delamare-Deboutteville & Massoud, 1964a: 80.
Specimens with transversal stripes and spots of dark pigment. Ant. IV with ten subsegments; Ant. III with 21 chaetae other than the sensory clubs, two of them as small sensilla in individual cavities; Ant. II with 15 chaetae, one of them as a small sensillum in cavity, four of the regular chaetae clearly longer than the others. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines absent. Trochanters I–III with 1,0,1 spines, respectively, trochanters I and III spines blunt; trochanter III with four regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and surpassing the tip of the unguis III. Female with a long subanal appendage (surpassing the ventral anal valves), spatulated, apically serrated on both edges. Dens ventral chaetotaxy formula from the apex to the base as: 2,2…1, dorsal chaetotaxy with 24 chaetae; mucronal notch prominent (adapted from
The genus Sminthurotheca Delamare-Deboutteville & Massoud, 1964 was erected based on a supposedly unique combination of Ant. III and large abdomen chaetotaxy. It was posteriorly synonymized with Sphyrotheca by
Specimens were found in gallery forests, in plant debris (
Angola, Congo (
Sphyrotheca mucroserratus Snider, 1978: 236.
Antennae, legs, and furca purplish, head purplish near the mouth, with purple bands between the antennae and on its vertex, trunk laterally with pale purple bands, posterior abdomen purplish. Ant. IV with nine or ten subsegments. Eyepatches with two interocular chaetae modified into strong spines each. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 absent. Trochanter III spine blunt, with five extra regular chaetae. Ungues with a single inner tooth, with tunica but lacking pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thin and reaching the tip of the unguis III. Large abdomen lacking capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), spatulated or acuminated, serrated at the tip. Manubrium with 7+7 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 14 chaetae; mucronal notch discrete (adapted from
The short description of
Specimens listed in the original description were found associated with Australian pine needles, leaf mold, and forest debris in Florida, USA (
Brazil, Mexico, and USA (
Sphyrotheca nepalica Yosii, 1966: 527.
Pale ground, body diffusely pigmented with brownish violet patches between the eyes and lateral sides of the large abdomen, antennae distally dark pigmented. Ant. IV with ten subsegments. Head vertex with a total of 11 large spines, three of them unpaired, including chaeta A1. Trochanters I–III with 1,1,1 spines, respectively, all blunt. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with or without the internal tooth; unguiculus III filament not reaching the tip of the unguis III. Large abdomen without capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, apically serrated on both edges. Manubrium with 6+6 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 13 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca nepalica is the only species of the genus with 11 spines on head vertex. Although the species fits Szeptyckitheca, especially due to the presence of the spines on trochanters I–III, its description is quite limited considering the current taxonomy of Symphypleona (see Table
Unknown.
Nepal (
Sphyrotheca peteri Palacios-Vargas, Vázquez & Cuéllar, 2003: 298–302, figs 1–3, Mexico, Quintana Roo, Reserva de la Biosfera de Sian Ka’na (orig. descr.).
Males pale bluish, with purple or blue spots on antennae, dorsal head, dorsal trunk and furca, females yellowish, with brown spots with the same distribution of males. Ant. IV with nine or ten subsegments, with some proximal chaetae capitate; Ant. II with 12 chaetae, two of them clearly longer than the others. Eyepatches with one interocular chaeta each. Head vertex with a total of 16 large and rough spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaetae near the spines present. Trochanters I–III with 1,1,1 spines respectively, trochanter II spine knobbed and III blunt; trochanter III with five regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and surpassing the tip of the unguis III. Large abdomen with capitate chaetae. Female with a long subanal appendage (surpassing the ventral anal valves), spatulated, apically serrated on both faces. Dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 17 chaetae; mucronal notch discrete (adapted from
Sphyrotheca peteri is herein transferred to Szeptyckitheca due to the presence of robust and somewhat erect spines on the head vertex and dorsal large abdomen, presence of three transversal rows of dental ventral chaetae and presence of spines on all trochanters, all features originally listed by
Specimens were found in low flooded jungle (
Mexico (
Sphyrotheca santiagoi
Yosii, 1959: 58;
Color pattern variable, usually mostly dark with a pale dorsum. Ant. IV with ten subsegments; Ant. III with at least 15 chaetae other than the sensory clubs; Ant. II with at least 15 chaetae, one of them clearly longer than the others; Ant. I with five chaetae. Eyepatches with one interocular somewhat spine-like chaeta each. Head vertex with a total of 14 or 16 spines, two of them unpaired; unpaired chaeta A1 absent. Trochanters I–III with 1,1,1 spines, respectively, all blunt; trochanter III with five regular chaetae other than the spine. Ungues without the inner tooth, with tunica and weak pseudonychia; unguiculus I with or without the internal tooth; unguiculus III filament thin and reaching the tip of the unguis III. Large abdomen without capitate mac. Female with a long subanal appendage, bidentate at the apex, smooth or apically serrated. Dens ventral chaetotaxy formula from the apex to the base as: 2,2…1, dorsal chaetotaxy with 13 chaetae; mucronal notch prominent (adapted from
Szeptyckitheca santiagoi is the sole species of the genus without the ungual inner tooth. However, the variability of color patterns reported by
Specimens were found in forest moss and litter, beach debris, up palms and in native gardens (
Australia, Papua New Guinea, Singapore, Solomon Islands (
Sphyrotheca spinimucronata
Itoh, 1993, in
White ground with diffuse purple pigment on anterior and dorsal head and dorsal and lateral trunk, antennae darker. Ant. IV with ~ 10 subsegments. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 present; secondarily reduced chaetae near the spines absent. Trochanter III spine blunt, with five extra regular chaetae. Ungues with a single inner tooth, with tunica and lacking pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen without capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, apically serrated on both edges. Manubrium with 7+7 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 17 chaetae; mucronal notch prominent (adapted from
Sphyrotheca spinimucronata is listed as a Szeptyckitheca species in
Specimens were found in a coniferous forest of Cryptomeria fortunei (
China (
Sphyrotheca vanderdrifti Delamare-Deboutteville & Massoud, 1964b: 64.
Yellowish ground color, with indistinct pigmented fields and purplish spots on the dorsum, antennae purplish. Ant. IV with ~ 10 subsegments; Ant. III with 19 chaetae other than the sensory clubs; Ant. II with 14 chaetae, two or three of them clearly longer than the others; Ant. I with six chaetae. Eyepatches with two interocular chaetae modified into strong spines each. Head vertex with a total of 16 large spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaeta near the spines present. Trochanters I–III with 1,1,1 spines, respectively, trochanter I spine acuminate and III capitate; trochanter III with four regular chaetae other than the spine. Ungues with a single inner tooth, with tunica and strong pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen lacking capitate mac. Female with a long subanal appendage (surpassing the ventral anal valves), acuminate, and apically or almost entirely serrated on its both edges. Dens ventral chaetotaxy formula from the apex to the base as: 2,1…1, dorsal chaetotaxy with 16 chaetae; mucronal notch discrete (adapted from
Specimens were found on marshy wood on sandy loam and shrubs on a ridge (
Suriname (
Holotype male on slide, Brazil, Piauí state, Altos municipality, “Floresta Nacional de Palmares” (5°3'12.53"S, 42°35'36.95"W), in sandy soil, Cerrado biome, 06/IV/2022, Mesquita C.P. col., pitfall traps. Paratypes on slides: one male, two females, one juvenile, with the same data as the holotype.
Ground color pinkish, with purple spots on head, dorso-anterior large abdomen, and dorsal small abdomen. Ant. IV with 11 subsegments, with five capitate chaetae; Ant. III with 19 chaetae other than the sensory clubs, two of them clearly longer than the others; Ant. II subdivided, with 13 chaetae, three of them clearly longer than the others; Ant. I with six chaetae. Eyepatches lacking interocular chaetae. Head vertex with a total of 14 large spines, two of them unpaired; unpaired chaeta A1 absent; secondarily reduced chaetae near the spines present. Trochanters I–III with 2,1,1 spines, respectively, trochanter I spines capitate, II and III spines blunt; trochanter III with five regular chaetae other than the spine. Ungues with one internal tooth, with tunica and weak pseudonychia; unguiculus I without the internal tooth; unguiculus III filament thin and reaching the tip of the unguis III. Large abdomen dorsally with 15+15 long capitate mac. Female with a short subanal appendage (not reaching the apex of the ventral anal valves), spoon-like, and apically serrated on both faces. Manubrium with 7+7 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 3…1, dorsal chaetotaxy with 17 chaetae; mucronal notch discrete.
Body (head + trunk) length of the type series ranging between 900 µm and 1400 µm, holotype with 900 µm, male average size = 900 µm, female average size = 1300 µm, entire type series average size = 1100 µm. Ground color pinkish, with purple spots on head, Ant. I–III, dorso-anterior large abdomen and dorsal small abdomen. Ant. IV, legs, and dens uniformly purplish (Fig.
Head (Figs
Trunk (Figs
Szeptyckitheca andrzeji sp. nov. A habitus in ethanol (dorsal view) B frontal head spine on papilla, red circle marks a secondarily reduced spine C large abdomen capitate mac D parafurcal area, red circle marks the neosminthuroid chaeta E bothriotrichum D, black arrows indicate small blunt accessory chaetae.
Szeptyckitheca andrzeji sp. nov. head A dorsal Ant. IV (white arrow points to chaeta present or absent) B dorsal Ant. III C dorsal Ant. II D dorsal Ant. I E anterior head chaetotaxy and eyes F labial and postlabial (ventral) chaetotaxy (right side) G maxillary outer lobe and sublobal plate (right side) H labial papillae and proximal chaetae alveoli (right side) I labrum J maxilla capitulum (right side) K mandibles apexes.
Abdominal appendages (Fig.
Legs (Figs
The species honors Dr. Andrzej Szeptycki for his important contributions to the taxonomy and systematics of springtails.
Specimens of Szeptyckitheca andrzeji sp. nov. were collected in the National Forest of Palmares, a small federal conservation unit, with a total area of 168.21 hectares, and altitudes ranging between 154 m and 250 m, located in Altos municipality, Piauí state, close to Teresina, the state’s capital (Fig.
Known distribution of Szeptyckitheca andrzeji sp. nov. and Szeptyckitheca cyanea sp. nov. in Brazil A National Forest of Palmares, in Altos municipality, Piauí state B Federal University of Rio Grande do Norte, in Natal city, Rio Grande do Norte state C National Forest of Nísia Floresta, in Nísia Floresta municipality, Rio Grande do Norte state. In A and C, the white line polygons delimit the sampled areas boundaries.
The climate in the region, according to the Köppen–Geiger climate classification system, is Aw, megathermic tropical with a long dry season and rainy summer, with high temperatures throughout the year (
Szeptyckitheca andrzeji sp. nov. is unique among the Neotropical taxa due to its reduced ventral dens chaetotaxy, with only four chaetae distributed in two transversal rows, following the formula from the apex to the basis: 3...1. All other Neotropical Szeptyckitheca species have three transversal lines of dental ventral chaetae. Also, this is the only species of the genus with a short subanal appendage, not reaching the apex of the ventral anal valves (Tables
Holotype male on slide, Brazil, Rio Grande do Norte state, Nísia Floresta municipality, “Floresta Nacional de Nísia Floresta” (6°5'9.132"S, 35°10'53.857"W), 02/VI/2022, Xavier M.D. col., pitfall traps. Paratypes on slides: one male and one female, with the same data as the holotype.
Two males and one female on slides, Brazil, Rio Grande do Norte state, Natal municipality, “Mata da CAERN – UFRN” (5°50'9.665"S, 35°12'12.953"W).
Specimens mostly bluish. Ant. IV with 11 subsegments, with at least six capitate chaetae; Ant. III with 21 chaetae other than the sensory clubs, including two peculiar small sensilla within cavities; Ant. II undivided, with 15 chaetae, four of them slightly longer than the others; Ant. I with six chaetae. Eyepatches with two small interocular chaetae. Head vertex with a total of 18 large spines, two of them unpaired; unpaired chaeta A1 present and regular (not spine-like); secondarily reduced chaetae near the spines absent. Trochanters I–III with 2,1,1 spines, respectively, trochanters I and II spines capitate, III blunt; trochanter III with five regular chaetae other than the spine. Ungues with a one inner tooth, with tunica and weak pseudonychia; unguiculus I with the internal tooth; unguiculus III filament thin and not reaching the tip of the unguis III. Large abdomen dorsally with ~ 26+26 long capitate mac. Female with a long subanal appendage (surpassing the apex of the ventral anal valves), slightly curved at the apex, acuminate, and apically serrated on its internal face. Manubrium with 7+7 dorsal chaetae; dens ventral chaetotaxy formula from the apex to the base as: 3,2…1, dorsal chaetotaxy with 16 chaetae; mucronal notch prominent.
Body (head + trunk) length of the type series ranging between 551 and 818 µm, holotype with 624 µm, males’ average size = 588 µm, females’ average size = 818 µm, entire type series’ average size = 664.5 µm. Specimens entirely dark bluish (Fig.
Head (Fig.
Szeptyckitheca cyanea sp. nov. head A dorsal Ant. IV (white arrows point to chaetae present or absent) B dorsal Ant. I–III, detail shows the ventral apical organ and the small sensilla in cavities on Ant. III C anterior head chaetotaxy and eyes D labial and postlabial (ventral) chaetotaxy (left side), white arrow points to chaeta present or absent E maxillary outer lobe and sublobal plate (left side) F labial papillae and proximal chaetae alveoli (left side) G labrum H mandibles apexes I maxilla capitulum (left side).
Trunk (Fig.
Abdominal appendages (Fig.
Legs (Figs
Szeptyckitheca cyanea sp. nov. legs A femur and tibiotarsus I (detail shows oval organs with internal small sensillum) B foot complex I C femur and tibiotarsus II (detail shows oval organs with internal small sensillum) D foot complex II E femur and tibiotarsus III (detail shows oval organs with internal small sensillum) F foot complex III.
The species was named after its color pattern; cyanea from Latin means dark blue.
Specimens of S. cyanea sp. nov. were found in two localities ~ 30 km apart in the Rio Grande do Norte state, Brazil: in the central campus of the Federal University of Rio Grande do Norte, Natal municipality, and the National Forest of Nísia Floresta, Nísia Floresta municipality (Fig.
Szeptyckitheca cyanea sp. nov. is the sole species of the genus with 18 spines on the head vertex. It is also the only Neotropical Szeptyckitheca with the frontal head A1 chaeta (see Table
1 | Head vertex with a set of 9 rough curved mac plus 4 large erect spines | S. koreana (Betsch & Weiner, 2009), comb. nov. (North Korea) |
– | Head vertex without rough curved mac, with 6 or more large erect spines | 2 |
2 | Head vertex with 15 large spines, ventral dens chaetotaxy formula as: 3,1…1, ungual pseudonychia absent | S. formosana (Yosii, 1965) (Taiwan) |
– | Head vertex with 6–14 or 16–18 large spines, ventral dens chaetotaxy formula otherwise, ungual pseudonychia usually present | 3 |
3 | Head vertex with 10 large spines, ungual tunica absent or at most vestigial, female subanal appendage very long, surpassing the dorsal anal valve | S. boneti (Denis, 1948) (Vietnam) |
– | Head vertex with 6 or 11–18 spines, ungual tunica present, female subanal appendage not reaching the apex of the dorsal anal valve | 4 |
4 | Head clypeal f line with 4+4 mac, ventral dens chaetotaxy formula as: 4,1…1, dorsal dens with 15 chaetae | S. kac Zeppelini, Lopes & Lima, 2018 (Brazil) |
– | Head clypeal f line without mac, ventral dens chaetotaxy formula otherwise, dorsal dens with 12–14 or 16–24 chaetae | 5 |
5 | Trochanter II without the spine, dorsal dens with 24 chaetae | S. machadoi (Delamare-Deboutteville & Massoud, 1964) (Angola, Congo) |
– | Trochanter II usually with the spine, dorsal dens with ≤ 20 chaetae | 6 |
6 | Head interantennal area with bifid chaetae, female anal valve with mps1 winged, dorsal dens with 20 chaetae | S. kesongensis Betsch & Weiner, 2009 (North Korea) |
– | Head interantennal area without bifid chaetae, female anal valve with mps1 not winged, dorsal dens with < 20 chaetae | 7 |
7 | Head vertex with 6 large spines, ventral dens chaetotaxy formula as: 4,2…1 |
S. coerulea ( |
– | Head vertex with 11 or more large spines, ventral dens chaetotaxy formula otherwise | 8 |
8 | Specimens pale, with diffused pigment and violet patches, head vertex with 11 large spines | S. nepalica (Yosii, 1966) (Nepal) |
– | Specimens color pattern otherwise, head vertex with 14–18 large spines | 9 |
9 | Specimens with lateral weak purple bands, posteriorly purple, dorsal dens with 14 chaetae | S. mucroserrata (Snider, 1978) (Brazil, Mexico, USA) |
– | Specimens color pattern otherwise, dorsal dens with 12–13 or 16–19 chaetae | 10 |
10 | Trochanter II without the spine, dorsal dens with 12 chaetae | S. karlarum (Palacios-Vargas, Vázquez & Cuéllar, 2003), comb. nov. (Mexico) |
– | Trochanter II with the spine, dorsal dens with > 12 chaetae | 11 |
11 | Ungues without the inner tooth, dorsal dens with 13 chaetae | S. santiagoi (Yosii, 1959) (Australia, Papua New Guinea, Singapore, Solomon Isl) |
– | Ungues with 1 inner tooth, dorsal dens 16–19 chaetae | 12 |
12 | Head frontal chaeta A1 present | 13 |
– | Head frontal chaetae A1 absent | 14 |
13 | Head vertex with 18 large spines, eyepatches with 2 interocular chaetae, ventral dens chaetotaxy formula as: 3,2…1, dorsal dens with 16 chaetae | S. cyanea Oliveira, Medeiros & Bellini, sp. nov. (Brazil) |
– | Head vertex with 16 large spines, eyepatches with 1 interocular chaeta, ventral dens chaetotaxy formula as: 2,2…1, dorsal dens with 19 chaetae | S. implicata (Hüther, 1967) (South Sudan) |
14 | Female subanal appendage short (not reaching the apex of the ventral anal valves), ventral dens chaetotaxy formula as: 3…1 | S. andrzeji Medeiros, Bellini & Weiner, sp. nov. (Brazil) |
– | Female subanal appendage long (surpassing the apex of the ventral anal valves), ventral dens chaetotaxy formula as: 3–2,2…1 | 15 |
15 | Head vertex with 14 large spines, trochanter I with 2 spines, unguiculus I without the internal tooth, mucronal notch prominent | S. bellingeri (Betsch, 1965) (Jamaica) |
– | Head vertex with 16 large spines, trochanter I with 1 spine, unguiculus I with the internal tooth, mucronal notch discrete | 16 |
16 | Ant. II with 14 chaetae, trochanter III spine capitate, with 4 regular chaetae, ventral dens chaetotaxy formula as: 3,2…1, dorsal dens with 17 chaetae | S. peteri (Palacios-Vargas, Vázquez & Cuéllar, 2003), comb. nov. (Mexico) |
– | Ant. II with 12 chaetae, trochanter III spine blunt, with 5 regular chaetae, ventral dens chaetotaxy formula as: 2,2…1, dorsal dens with 16 chaetae | S. vanderdrifti (Delamare-Deboutteville & Massoud, 1964) (Suriname) |
*We did not include S. spinimucronata as we considered it a species inquirenda. Further details on the species are listed in its diagnosis.
The boundaries between Szeptyckitheca and Sphyrotheca are quite narrow (
After the comparison of both new species described here, we observed other features which may be useful to diagnose Szeptyckitheca taxa, such as the shape of the maxilla capitulum (globose vs elongate), the shape of E chaetae on frontal head, the presence of an extra basal appendix on proximal dens and the number, and homology of clypeal unpaired chaetae. These features were not used in this study to separate the species due to the lack of information or uncertainties about such data in most (or all) previously described species. Nevertheless, with the expansion of the knowledge of the genus they may be useful to identify closely related taxa.
After our study, there are now 18 species assigned to Szeptyckitheca. Even so, many of them are in need of revision or full redescription.
We would like to thank the anonymous reviewers for carefully revising the manuscript and providing ideas to improve it.
The authors have declared that no competing interests exist.
Ethical review and approval were waived for this study due to Brazilian laws which do not require permission from an institutional ethics committee on the use of animals for taxonomical studies with microarthropods.
This research was funded by the National Council for Scientific and Technological Development (CNPq), grant number 309114/2021-7 (BCB project), Institutional Scientific Initiation Scholarship Program (PIBIC/CNPq) (MFDO scholarship) and the Coordination for the Improvement of Higher Education Personnel (CAPES), grant number 001 (GDSM scholarship).
Conceptualization: BCB, GDSM. Data curation: BCB, MFDO, GDSM. Formal analysis: MFDO, GDSM. Project administration: BCB. Resources: BCB. Software: RCN, GDSM. Supervision: BCB, WMW. Validation: GDSM, BCB. Visualization: BCB, WMW, GDSM. Writing - original draft: BCB, RCN. Writing - review and editing: GDSM, RCN, BCB, MFDO, WMW.
Bruno Cavalcante Bellini https://orcid.org/0000-0001-7881-9436
Wanda Maria Weiner https://orcid.org/0000-0002-7257-3671
Rudy Camilo Nunes https://orcid.org/0000-0002-3140-9146
Gleyce Da Silva Medeiros https://orcid.org/0000-0001-9839-2345
All of the data that support the findings of this study are available in the main text.