Table 1.
Recent scorpion families, genera and species without median and/or lateral ocelli.
Table 2.
Lateral ocellus counts in Recent scorpion families, according to the literature.
Table 3.
Number of genera, species and individuals examined for study of lateral ocelli in Recent scorpion families.
Figure 1.
Six-ocellus model with four accessory ocelli, indicated in black (A), and nine general patterns with fewer than six ocelli (B–J), indicating most common positions and relative sizes (AALMa <ALMa <MLMa> PLMa> PDMi>ADMi; PDMi> PLMi; PLMi> APLMi1; PLMi = APLMi2; PLMi> APLMi3) of lateral ocelli in Recent scorpions.
B. Type 5, shaded areas represent positional variation in ADMi and PDMi. C. Type 4A. D. Type 4B. E. Type 4C. F. Type 3A. G. Type 3B. H. Type 2A. I. Type 2B. J. Type 1. Abbreviations: accessory anterolateral major ocellus (AALMa); anterolateral major ocellus (ALMa); mediolateral major ocellus (MLMa); posterolateral major ocellus (PLMa); posterolateral minor ocellus (PLMi); posterodorsal minor ocellus (PDMi); anterodorsal minor ocellus (ADMi); three accessory posterolateral minor ocelli (APLMi1, APLMi2, APLMi3).
Figure 2.
Lateral ocelli of Recent scorpions (family Buthidae C.L. Koch, 1837).
A. Androctonus finitimus (Pocock, 1897), subad. ♂ (AMNH), Type 5. B. Tityopsis inexpectata (Moreno, 1940), ♀ (AMNH), abnormal six-ocellus condition with AALMa on sinistral side. C. Buthacus macrocentrus (Ehrenberg, 1828), subad. ♀ (AMNH), Type 5. D. Centruroides vittatus (Say, 1821), ♀ (AMNH), Type 5. E. Orthochirus scrobiculosus (Grube, 1873), ♀ (AMNH), Type 5. F. Razianus zarudnyi (Birula, 1903), ♀ (AMNH), Type 4C. G. Tityus bahiensis eickstedtae Lourenço, 1982, juv. (AMNH [LP 1596]), Type 5. H. Alayotityus nanus Armas, 1973, ♂ (AMNH [LP 1768]), Type 4A. I. Microtityus consuelo Armas & Marcano Fondeur, 1987, ♀ (AMNH [LP 3281]), Type 4A. J. Isometrus maculatus (DeGeer, 1778), ♂ (AMNH [LP 1798]), Type 5. K. Zabius fuscus Thorell, 1876, subad. ♀ (AMNH), Type 4C. L. Akentrobuthus atakora Vignoli & Prendini, 2008, holotype ♀ (AMNH [LP 8334]), Type 2A. M. Uroplectes carinatus (Pocock, 1890), ♂ (AMNH), Type 5. N. Afroisometrus minshullae (FitzPatrick, 1994), ♀ (AMNH [LP 7875]), Type 3A. O. Pseudolychas ochraceus (Hirst, 1911), ♀ (AMNH), Type 5. P. Lissothus bernardi Vachon, 1948, ♀ (MNHN RS 3420), Type 4C. Q. Karasbergia methueni Hewitt, 1913, ♂ (AMNH [LP 8225]), Type 3A. R. Pseudolychas ochraceus (Hirst, 1911), ♀ (AMNH), Type 4C. Abbreviations: AALMa (accessory anterolateral major ocellus); ADMi (anterodorsal minor ocellus); ALMa (anterolateral major ocellus); e (eyespot); MLMa (mediolateral major ocellus); PDMi (posterodorsal minor ocellus); PLMa (posterolateral major ocellus). Scale bars = 0.1 mm.
Figure 3.
Lateral ocelli of Recent scorpions (families Chactidae Pocock, 1893; Chaerilidae, Pocock, 1893; Euscorpiidae, Laurie, 1896; Iuridae Thorell, 1876; Pseudochactidae Gromov, 1998; Scorpiopidae Kraepelin, 1905).
A. Chaerilus variegatus Simon, 1877, ♀ (AMNH), abnormal three-ocellus condition with PLMi. B. C. variegatus, ♂ (AMNH [LP 6390]), abnormal three-ocellus condition with ALMa. C. C. variegatus, ♂ (AMNH [LP 6389]), Type 2A. D, E. Chaerilus chapmani Vachon & Lourenço, 1985, ♀ (AMNH), Type 2A on sinistral side (D), abnormal one-ocellus condition without MLMa (Type 1) on dextral side (E). F. Pseudochactas ovchinnikovi Gromov, 1998, subad. ♂ (AMNH), Type 1. G. Troglocormus ciego Francke, 1981, holotype ♂ (AMNH), Type 4C. H. Protoiurus kraepelini (von Ubisch, 1922), ♀ (AMNH), Type 4C. I. Calchas birulai Fet et al., 2009, ♀ (AMNH), Type 4C. J. Megacormus gertschi Díaz Najera, 1966, juv. ♂ (AMNH [LP 6474]), Type 4C. K. Euscorpius italicus (Herbst, 1800), ♂ (AMNH [LP 10297]), Type 3B. L. Troglocormus willis Francke, 1981, ♀ (AMNH), Type 3B. M. Chactopsis insignis Kraepelin, 1912, ♀ (AMNH [LP 8420]), Type 4C. N. Parascorpiops montana Banks, 1928, lectotype ♂ (MCZ), Type 3A. O. P. montana, paralectotype ♀ (MCZ), Type 2A. P. Alloscorpiops sp., ♀ (AMNH [LP 11279]), Type 4B. Q. Scorpiops feti Kovařík, 2000, ♀ (MCZ), Type 4B. R. Euscorpiops kaftani (Kovařík, 1993), juv. ♀ (AMNH [LP 11371]), Type 4B. Abbreviations: ADMi (anterodorsal minor ocellus); ALMa (anterolateral major ocellus); e (eyespot); MLMa (mediolateral major ocellus); PDMi (posterodorsal minor ocellus); PLMa (posterolateral major ocellus); PLMi (posterolateral minor ocellus). Scale bars = 0.1 mm.
Figure 4.
Lateral ocelli of Recent scorpions (families Chactidae Pocock, 1893; Iuridae Thorell, 1876; Troglotayosicidae Lourenço, 1998).
A. Megachactops kuemoi Ochoa et al., 2013, juv. paratype (AMNH [LP 9244]), Type 4C. B. Chactopsoides anduzei (González-Sponga, 1982), ♀ (AMNH), Type 4C. C. Taurepania porosus (Pocock, 1900), ♀ (AMNH [LP 5513]), Type 4C. D. Hadrurochactas machadoi González-Sponga, 1993, subad. ♀ (AMNH), Type 4C. E. Broteochactas nitidus Pocock, 1893, ♀ (AMNH [LP 1511]), Type 4C. F. Teuthraustes glaber Kraepelin, 1912, ♀ (AMNH), Type 3A. G. Vachoniochactas lasallei (González-Sponga, 1978), subad. ♂ (AMNH [LP 10000]), Type 4C. H. Chactas raymondhansi Francke & Boos, 1986, subad. ♀ (AMNH [LP 1586]), Type 2A. I. Troglotayosicus humiculum Botero-Trujillo & Francke, 2009, ♀ (AMNH), Type 2B. J. Brotheas granulatus Simon, 1877, ♂ (AMNH [LP 3656]), Type 4C. K, L. Neochactas delicatus (Karsch, 1879), ♀ (AMNH [LP 3441]), Type 4C (K), Type 3A (L). M, N. Anuroctonus phaiodactylus (Wood, 1863), ♂ (AMNH), Type 4C, sinistral side (M), dextral side (N). O. Hoffmannihadrurus gertschi (Soleglad, 1976), paratype ♀ (AMNH), Type 4C. P. Caraboctonus keyserlingii Pocock, 1893, ♂ (AMNH), Type 4C. Q. Hadruroides charcasus (Karsch, 1879), ♂ (AMNH), Type 4C. R. Hadrurus arizonensis Ewing, 1928, ♂ (IBUNAM), Type 3A. Abbreviations: ADMi (anterodorsal minor ocellus); e (eyespot); MLMa (mediolateral major ocellus); PDMi (posterodorsal minor ocellus); PLMa (posterolateral major ocellus). Scale bars = 0.1 mm.
Figure 5.
Lateral ocelli of Recent scorpions (families Bothriuridae Simon, 1880; Chactidae Pocock, 1893; Superstitioniidae Stahnke, 1940; Vaejovidae Thorell, 1876).
A. Superstitionia donensis Stahnke, 1940, ♀ (AMNH), Type 4C. B. Nullibrotheas allenii (Wood, 1863), ♀ (AMNH) Type 3A. C. Uroctonus mordax Thorell, 1876, ♂ (AMNH), Type 3A. D. Uroctonus mordax pluridens Hjelle, 1972, ♀ (AMNH [LP 2686]), Type 3A. E, F. U. m. pluridens, ♂ (AMNH [LP 2686]), Type 4C, dextral side (E), sinistral side (F). G. Uroctonites giulianii Williams & Savary, 1991, ♂ (AMNH), Type 3A. H. Paruroctonus surensis Williams & Haradon, 1980, ♂ (AMNH [LP 3139]), Type 3A. I. Smeringurus grandis (Williams, 1970), juv. ♀ (AMNH [LP 4462]), Type 3A. J. Vaejovis carolinianus (Beauvois, 1805), ♀ (AMNH [LP 8500]), Type 3A. K. Thorellius intrepidus (Thorell, 1876), ♀ (AMNH [LP 2022]), Type 3A. L. Stahnkeus subtilimanus (Soleglad, 1972), ♀ (AMNH), Type 3A. M. Bothriurus bonariensis (C.L. Koch, 1842), ♀ (AMNH), Type 3A. N. Timogenes mapuche Maury, 1975, ♂ (AMNH [LP 4312]), Type 3A. O. Vachonia martinezi Abalos, 1954, juv. ♀ (AMNH [LP 2441]), Type 2B. P. Centromachetes sp., ♂ (AMNH), abnormal four-ocellus condition with ADMi (Type 4C) on dextral side. Q. Lisposoma elegans Lawrence, 1928, ♂ (AMNH), abnormal four-ocellus condition with ALMa on dextral side. R. Lisposoma josehermana Lamoral, 1979, ♂ (AMNH), Type 3A. Abbreviations: ADMi (anterodorsal minor ocellus); ALMa (anterolateral major ocellus); e (eyespot); MLMa (mediolateral major ocellus); PDMi (posterodorsal minor ocellus); PLMa (posterolateral major ocellus). Scale bars = 0.1 mm.
Figure 6.
Lateral ocelli of Recent scorpions (families Diplocentridae Karsch, 1880; Hemiscorpiidae Pocock, 1893; Heteroscorpionidae Kraepelin, 1905; Hormuridae Laurie, 1896; Scorpionidae Latreille, 1802; Urodacidae Pocock, 1893).
Hemiscorpius lepturus Peters, 1861, ♂ (AMNH [LP 11080]), Type 3A. B. Heteroscorpion magnus Lourenço & Goodman, 2002, paratype ♂ (FMNH), Type 2B. C. Urodacus manicatus (Thorell, 1876), ♀ (AMNH), Type 2B. D. Cheloctonus jonesii Pocock, 1892, ♀ (AMNH), abnormal four-ocellus condition with PLMi (Type 4B) on dextral side. E. Cheloctonus jonesii Pocock, 1892, ♀ (AMNH), Type 3A. F. Hormiops davidovi Fage, 1933, ♀ (AMNH), Type 2B. G. Diplocentrus rectimanus Karsch, 1880, ♀ (AMNH [LP 2032]), abnormal four-ocellus condition with APLMi2 on dextral side. H. Nebo hierichonticus (Simon, 1872), ♂ (AMNH), Type 3A. I. Liocheles australasiae (Fabricius, 1775), ♀ (AMNH), Type 3A. J. Tarsoporosus macuira Teruel & Roncallo, 2010, ♂ (AMNH), Type 4B. K. Oiclus purvesii (Becker, 1880), ♀ (AMNH [LP 9037]), Type 2B. L. Pandinus gregoryi (Pocock, 1896), juv. ♂ (AMNH), Type 3A. M. Opistophthalmus jenseni (Lamoral, 1972), ♂ (AMNH), abnormal seven-ocellus condition with MLMa, PLMa, PLMi, PDMi, APLMi1, APLMi2 and APLMi3. N, O. O. jenseni, ♂ (AMNH [AH 4039]), abnormal five-ocellus condition with MLMa, PLMa, PDMi, PLMi and APLMi2 on sinistral side (N) and MLMa, PLMa, PDMi, PLMi and ADMi on dextral side (O). P. Opistophthalmus sp., ♂ (AMNH), Type 4B. Q. Scorpio maurus palmatus (Ehrenberg, 1828), ♂ (AMNH), Type 4B. R. Scorpio maurus palmatus (Ehrenberg, 1828), ♂ (AMNH), Type 3A. Abbreviations: anterodorsal minor ocellus (ADMi); APLMi1, APLMi2, APLMi3 (accessory posterolateral minor ocelli); MLMa (mediolateral major ocellus); PDMi (posterodorsal minor ocellus); PLMa (posterolateral major ocellus); PLMi (posterolateral minor ocellus). Scale bars = 0.1 mm.
Table 4.
General patterns (Types 5, 4A, 4B, 4C, 3A, 3B, 2A, 2B, 1) of lateral ocelli in Recent scorpion families and genera.
Table 5.
Recent scorpion families and genera in which an eyespot is present, including observations from the literature [27]–[30].