Systematics and biogeography of Salina MacGillivray (Collembola: Entomobryoidea), with emphasis on the species groups in the New World

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Nine new species of the collembolan genus Salina MacGillivray from South America are described and illustrated. Two Neotropical species were recorded for the first time from Brazil: S. dedoris Mari-Mutt and S. tristani Denis. Salina was previously known to occur in three Brazilian states, and this is now updated to include 19 states with 12 recorded species. A new proposal of morphological character description and illustration, and an identification key for the celebensis group are provided. A hypothesis for the phylogenetic relationships among 34 species of Salina (about 50% of the 72 described species) allowed three main pursuits: (a) a reevaluation of Salina species groups; (b) the first explicit interpretation of how morphological characters of these springtails may have changed during the course of the diversification of the taxon; and (c) an evaluation of the historical biogeographic connections of Salina, with an emphasis on the celebensis group distribution to the New World.

Insect Systematics & Evolution

An International Journal of Systematic Entomology

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Figures

  • Symbols of chaetae used in chaetotaxy of Salina spp.

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  • (A–B). Phylogenetic relationships among 34 species of Salina (as shown in Figure 20) and photographs illustrating color pattern diversity comprised. Color circles indicate biogeographical regions of species: Nearctic (blue circle), Neotropical (green), Afrotropical (brown), Madagascan (red), Oriental (yellow), and Australian (pink). “For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.”

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  • Salina serrana sp.n., habitus and tergal structures: (A) lateral habitus; (B), cuticle and ecdysial line; (C), pseudopore; (D), mac apically clubed; (E), mac apically acuminate; (F), mic apically branched; (G), bothriotrichum of Abd II (basal region); (H), S-microchaeta (ms) of Th II, Abd I and III; I, sens type I of Th II–III; (J), sens type II of Abd III; (K), sens type III of Abd V; (L), sens type IV of Abd IV; (M), sens type V (bs) present between T2 and T4 bothriotricha of Abd IV.

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  • Habitus of Salina spp. (dorsal view): (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n. (distal half of Ant IV missing); (I), S. unisetosa sp.n.; (J), S. dedoris from Roraima state, Brazil; (K), S. tristani from Amazonas state, Brazil. Scale bars: 0.5 mm.

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  • Salina serrana sp.n., antenna and head structures: (A), dorsal head, arrow indicates the “sutural series”; (B), lent of the D eye on right side (lateral view), yellow color indicates the differentiated cuticle; (C), dorsum of the Ant I base and anterior head, arrow indicates serrated mic of antennal series; (D), serrated An2a mic; E, apex of Ant IV, apical bulb not visible; (F), median part of Ant IV, arrow indicates annulations; (G), apex of Ant III, green marked indicates apical organ, orange marked indicates the ventro-external microsens, red marked indicates guard sensillae, and blue marked indicates the long finger-shaped sens. “For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.”

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  • Salina serrana sp.n., anterior head and mouthparts: (A), labral papillae, maxillary lobes, prelabral and labral chaetotaxy; (B), Chaeta a1 of labrum; (C), labral papillae; (D), maxillary outer lobe and sublobal plate appendages; (E), labial palp (a-e = labial papillae, h = hypostomal papilla) and labial basomedian (a1-2 smooth chaetae, M and E ciliate chaetae) and basolateral fields.

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  • New proposal of tergal homology for the celebensis group: (A), Abd III; (B), Abd IV, arrow indicates sens rarely present, arrow two ends indicates variable positions of B3 mac; (C), Abd V.

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  • Genital plate and furcula of Salina spp.: (A–B), Genital plate of Salina serrana sp.n. (ventral view): (A), ♂; (B), ♀. (C), Distal manubrium and proximal dens of S. brasiliana sp.n. (lateral view), arrow indicates chaeta absent in manubrial plate of some species. (D–I), distal dens and mucro (inner view): (D), S. brasiliana sp.n.; (E), S. bellinii sp.n.; (F), S. hamadae sp.n.; (G), S. maculipenis sp.n.; (H), S. peruana sp.n.; (I), S. serrana sp.n.

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  • Salina serrana sp.n., abdominal appendages: (A), collophore (ventral view); (B), tenaculum (ventral view); (C), furcula (lateral view); (D), distal dens and mucro (internal view); (E), distal dens and mucro (posterior view).

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  • Head structures of Salina spp.: (A–I), Ant I dorsal chaetotaxy (right): (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n.; (I), S. unisetosa sp.n., arrow indicates mac present or absent. (J–K), general structures of anterior and ventral head: (J), labral papillae, prelabral and labral chaetotaxy; (K), maxillary outer lobe and sublobal plate; (L), labial palp and proximal chaetae; (M), labial basolateral and basomedian fields.

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  • New proposal for tergal homology for the celebensis group (morphological terminology adapted from Szeptycki, 1979). (A), head; (B), Th II; (C), Th III; (D), Abd I; (E), Abd II.

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  • Head ventral chaetotaxy of Salina spp.: (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n.; (I), S. unisetosa sp.n.

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  • Legs structures of Salina spp.: (A–I), trochanteral organ (right): (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n.; (I), S. unisetosa sp.n. (J–K), distal tibiotarsus and foot complex (posterior side): (J), fore leg; (K), hind leg.

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  • Salina serrana sp.n., legs structures: (A), distal tibiotarsus and foot complex I (anterior side); (B), distal tibiotarsus and empodial complex III (posterior side); (C), connection of empodial complex III (anterior side), arrow indicates paired external teeth; (D), connection of empodial complex III (posterior side), red arrow indicates posterior pretarsal chaeta, white arrow indicates inner smooth chaetae; (E), external side of distal tibiotarsus and unguis III; (F), tenent-hair III (laterodorsal view); (G), apical region of tenent-hair III. “For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.”

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  • Th II–III dorsal chaetotaxy of Salina spp.: (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n.; (I), S. unisetosa sp.n.

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  • Abd I dorsal chaetotaxy of Salina spp.: (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. peruana sp.n.; (G), S. serrana sp.n.; (H), S. tocantinensis sp.n.; (I), S. unisetosa sp.n.

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  • Abd III–IV dorsal chaetotaxy of Salina spp.: (A–C), Abd III: (A), Salina brasiliana sp.n.; (B), S. hamadae sp.n.; (C), S. serrana sp.n. (D–I), Abd IV: (D), S. brasiliana sp.n.; (E), S. hamadae sp.n.; (F), S. serrana sp.n.; (G), S. bicolor sp.n.; (H), S. maculipenis sp.n.; (I), S. peruana sp.n.

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  • Abd V dorsal chaetotaxy of Salina spp.: (A), Salina brasiliana sp.n.; (B), S. bellinii sp.n.; (C), S. bicolor sp.n.; (D), S. hamadae sp.n.; (E), S. maculipenis sp.n.; (F), S. serrana sp.n.; (G), S. tocantinensis sp.n.

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  • Collophore chaetotaxy of Salina spp.: (A–B), Salina brasiliana sp.n.: (A), anterior side; (B), lateral and posterior side. (C–D), S. bellinii sp.n.: (C), anterior side; (D), lateral and posterior side. (E), complete chaetotaxy (lateral view) of S. bicolor sp.n. (F–G), S. hamadae sp.n.: (F), anterior side; (G), lateral and posterior side.

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  • Collophore chaetotaxy of Salina spp.: (A–B), Salina maculipenis sp.n.: (A), anterior side; (B), lateral and posterior side. (C–E), complete chaetotaxy (lateral view): (C), S. peruana sp.n.; (D), S. serrana sp.n.; (E), S. unisetosa sp.n.; (F–G), S. tocantinensis sp.n.: (F), anterior side; (G), lateral and posterior side.

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  • Phylogenetic relationships among species of Salina (54 steps, CI= 50, RI=76): (A). One of the 12 most parsimonious trees, grey squares showing value of of the Bremer support for the branches, character state transformations are depicted as black circles for unequivocal changes and white circles for reversed states or with multiple changes. (B). Strict consensus of the 12 most parsimonious trees, grey shades indicates the limits of the species groups beta, celebensis, and representatives of yosii and borneensis groups comprised in clade A; two unnamed species nested within the celebensis group cannot be placed in any of the traditionally recognized groups. (C). Some morphological characters analyzed for this study: 1-3, dens and mucro of Akabosia matsudoensis, S. maculiflora and S. peruana sp.n., respectively; 4, head dorsal chaetotaxy (right side) of Salina sp. 4 (borneensis group); 5-7, Abd II chaetotaxy (right side) of S. beta (beta group), S. hamadae sp.n. (celebensis group) and Salina sp. 4 (borneensis group), respectively; 8, Abd IV chaetotaxy (right side) of Salina sp. 4 (borneensis group).

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  • Geographic distribution of lineages of Salina from the the Neotropic and Nearctic regions included in this study: (A–C), celebensis group: (A), S. brasiliana; (B), S. bellinii sp.n. (red), S. dedoris (green), S. tristani (yellow), S. zhangi (blue). (C), (1) S. peruana sp.n. (pink) and S. unisetosa sp.n. (grey); (2) S. serrana sp.n. (yellow) and S. hamadae sp.n. (purple); (3) S. panamae (orange) and S. maculipenis sp.n. (black); S. bicolor sp.n. (red), S. tocantinensis sp.n. (green), S. banksi (blue). (D), beta group: S. maculiflora (orange), S. colombiana (red), S. wolcotti (blue), S. bidentata (green) and S. beta (black). “For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.”

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