Genus Charinus Simon, 1892

Figs 2–102; Tables 1–7

Charinus Simon, 1892: 48

(type species: Phrynus australianus L. Koch, 1867, by original designation).

Enantiosarax Mello-Leitão, 1931: 55 (synonymized by Quintero 1983: 46) (type species: Enantiosarax schirchii Mello-Leitão, 1931, by monotypy).

Oligacanthophrynus Caporiacco, 1947: 21 (synonymized by Weygoldt 1998a: 72) (type species: Oligacanthophrynus guianensis Caporiacco, 1947, by original designation).

Speleophrynus Ravelo, 1975: 78–79 (synonymized by Quintero 1983: 27) (type species: Speleophrynus tronchonii Ravelo, 1975, by original designation).

Tricharinus Quintero, 1986: 206 (synonymized by Weygoldt 2000a: 25) (type species: Tricharinus guianensis Quintero, 1986, by original designation).

Charinus – Kraepelin 1895: 46; 1899: 248–249. — Gravely 1915b: 442. — Mello-Leitão 1931: 54. — Werner 1935: 471. — Weygoldt 1999b: 48; 2000a: 23–24. — Armas & Pérez 2001: 62. — Harvey 2003: 3–4. — Armas 2006b: 225; 2014: 31.

Oligacanthophrynus – Caporiacco 1948: 616.

Type species

Phrynus australianus L. Koch, 1867, by original designation.

Diagnosis

Charinus may be distinguished from the other two genera of Charinidae by the following combination of characters: lateral eyes situated at least three times diameter of one ocellus from carapace lateral margin, with seta posterior to lateral ocular triad (Fig. 37A–B); setiferous tubercles situated far apart on dorsal carina of pedipalp coxa; female gonopod cushion-like (Figs 5C–D, 7C–D), finger-like or suckerlike (Figs 5A–B, 7A–B); posterior margin of female genital operculum straight or rounded (Fig. 17); area of genital operculum between gonopods and posterior margin smooth in dorsal view (Fig. 17).

Unlike Charinus, in which the area of genital operculum between the gonopods and posterior margin is smooth, and the posterior margin curved dorsally, the area has denticulate projections in Sarax (Figs 106D, 113D), and the posterior margin is linear in Weygoldtia .

Etymology

Unknown, but likely to be the diminutive of the name Charon .

Description

CARAPACE. Frontal process well developed, much wider than long, with blunt, rebordered or acute apex (Figs 42C, 54C). Carapace dorsoventrally compressed, wider than long. Median eyes and ocular tubercle or, when tubercle absent, anterior depression, leading to narrow medial sulcus that extends around posterior area of pair of lateral humps situated behind lateral eyespots (Fig. 35A). Anterior margin rounded with fine setae and corners slightly depressed; with 6–10 small setae, two medial setae usually directly anterior to median ocular tubercle (Fig. 37A). Carina originating at anterolateral margin, widening posteriorly from coxa I and onwards, widest dorsal to coxae III and IV, and reaching posterior margin. Many tiny punctations, more abundant in anterior area (Fig. 33A), arranged in lines and spots, irradiating from fovea and interspersed with glabrous areas. Three pairs of deep sulci and very deep, rectangular or triangular fovea (corners form starting point of second and third sulci). First pair of sulci situated slightly posterior to lateral boss and not reaching midline. Four pairs of lateral depressions (first placed over first pair of sulci). Lateral eyes well developed, or reduced to small, whitish spot, without cornea or clearly defined lens (only small roundish knob), rarely absent.

STERNUM. Four-segmented, all platelets sclerotized and convex (Fig. 31B). Tritosternum with round base, projected anteriorly into long or small, blunt tubercle, with two apical setae and two basal setae (Fig. 24B). Medial platelet (tetrasternum) rounded, convex, with two setae and few setulae; formed by single (Fig. 30B) or paired (Fig. 24B) platelets. Third platelet (pentasternum) also rounded and convex, but smaller, with shorter setae than second platelet; formed by single (Fig. 30B) or paired (Fig. 24B) platelets. Metasternum paired in anterior half, with anterior setae in membranous region followed by two or three setae in sclerotized area, arranged in longitudinal row from unsclerotized to sclerotized region; distal border with small elevation bearing 2–8 large setae. Sternites separated from each other by twice diameter of medial platelet.

OPISTHOSOMA. Oblong, with almost indistinguishable punctations, finer than on carapace (Fig. 30A).

GENITALIA. Female gonopods rounded or oval,sucker-like (barrel shaped), with rounded opening(Fig. 65), finger-like (in C. africanus Hansen, 1921 only) or cushion-like, with wide atrial opening or covered with claw-like projection (Fig. 22). Male gonopods slightly wider than long, soft, only posterolateral margin of dorsal lobes and basal sclerite of lateral lobes sclerotized; median lobes long and thin, lamellar, with wide, rounded tip, almost reaching apex of lateral lobes (Fig. 25).

CHELICERAE. Cheliceral furrow with four prolateral teeth (Fig. 10), distal tooth bifid. Fourth tooth twice as long as others and much stouter. Tooth length (from tip to base) IV>Ia>Ib =II>III. Claw with 5–13 denticles (Fig. 11A–B, E–F).

PEDIPALPS. Coxa with dorsal carina rounded in shape; 2–5 equidistant setae along prolateral margin, and 0–6 setae encircled by round carina. Trochanter with large distal, spiniform, ventral apophysis, bearing many prominent setae, and with blunt apex pointed anteriorly, and two subequal spines, one in median third and other at distal apex of prolateral surface (Fig. 2). Femur with 2–7 dorsal spines in primary series, decreasing in size, sometimes with secondary row of spines (Fig. 3A–D); femur dorsal surface with row of 2–4 setiferous tubercles between spine 1 and proximal margin of segment; 2–6 ventral spines in primary series, sometimes with secondary row of spines; setiferous tubercle or spine between spine ventral 1 and proximal margin, aligned with primary series or displaced dorsally. Patella with 3–6 dorsal spines in primary series, decreasing in lengths (Fig. 4A–D); small setal tubercle between spine I and distal margin of patella; 2–6 ventral spines, distal larger; 1–4 setiferous tubercles or 1–2 spines between ventral spine I and distal margin. Tibia with two dorsal spines, distal at end of proximal half, proximal in proximal third (Fig. 30); ventral spine in distal half, around two-thirds length of tibia, and 2 or 3 setal tubercles proximal to spine; row of 2–5 long, thin setae (longer than others in vicinity). Tarsus with 1–3 curved spines in proximal half (Figs 13, 52D); cleaning organ about half article length of tarsus. Claw (apotele) with long, acute, curved apex.

LEGS. All very setose. Femur length on leg I>III>IV>II; ventral corner of prolateral surface of leg II–IV femora projecting into distinct spiniform process. Tibia of leg I with 16–25 articles (up to 47 in regenerated legs); distal articles of tibia each with two small trichobothria, one on dorsal and one on prolateral side of article; one trichobothrium on second, third and fourth (from distal to proximal) articles, close to distal border, all situated dorsally; no trichobothria on other articles. Leg I tarsus (basitarsus + distitarsus) with 23–43 articles (up to 62 in regenerated legs); apical article with claw, tarsal organ and rod sensilla (Fig. 16). Leg IV basitibia with 2–4 pseudo-articles, one trichobothrium on last pseudo-article. Leg IV distitibia with one basal, two median and 13–18 distal trichobothria. Leg IV basitibia–distitibia length BT1>DT>BT3=BT4>BT2. Leg IV basitarsus/distitarsus ratio 7/4, distitarsus tetramerous.

Included taxa

Charinus acaraje Pinto-da-Rocha, Machado & Weygoldt, 2002; Charinus acosta (Quintero, 1983); Charinus africanus Hansen, 1921; Charinus aguayoi Moyá-Guzmán, 2009; Charinus alagoanus sp. nov.; Charinus apiaca sp. nov.; Charinus asturius Pinto-da-Rocha, Machado & Weygoldt, 2002; Charinus australianus (L. Koch, 1867); Charinus bahoruco Teruel, 2016; Charinus belizensis Miranda, Giupponi & Wizen, 2016; Charinus bichuetteae Giupponi & Miranda, 2016; Charinus bonaldoi Giupponi & Miranda, 2016; Charinus bordoni (Ravelo, 1977); Charinus brasilianus Weygoldt, 1972; Charinus brescoviti Giupponi & Miranda, 2016; Charinus bromeliaea Jocqué & Giupponi, 2012; Charinus bruneti Teruel & Questel, 2011; Charinus caatingae Vasconcelos & Ferreira, 2016; Charinus camachoi (González-Sponga, 1998); Charinus carajas Giupponi & Miranda, 2016; Charinus caribensis (Quintero, 1986); Charinus carinae sp. nov.; Charinus carioca sp. nov.; Charinus carvalhoi sp. nov.; Charinus cavernicolus Weygoldt, 2006, new rank; Charinus cearensis sp. nov.; Charinus centralis Armas & Ávila Calvo, 2000; Charinus cubensis (Quintero, 1983); Charinus decu (Quintero, 1983); Charinus desirade Teruel & Questel, 2015; Charinus diamantinus sp. nov. Charinus diblemma Simon, 1936, nomen dubium; Charinus dominicanus Armas & Pérez, 2001; Charinus elegans Weygoldt, 2006, stat. nov.; Charinus eleonorae Baptista & Giupponi, 2003; Charinus euclidesi sp. nov.; Charinus fagei Weygoldt,1972; Charinus ferreus Giupponi &Miranda, 2016; Charinus gertschi Goodnight& Goodnight, 1946; Charinus goitaca sp. nov.; Charinus guayaquil sp. nov.; Charinus guianensis (Caporiacco, 1947), nomen dubium; Charinus guto Giupponi & Miranda, 2016; Charinus imperialis sp. nov.; Charinus insularis Banks, 1902; Charinus iuiu Vasconcelos & Ferreira, 2016; Charinus jeanneli Simon, 1936, nomen dubium; Charinus jibaossu Vasconcelos, Giupponi & Ferreira, 2014; Charinus kakum Harms, 2018; Charinus koepckei Weygoldt, 1972; Charinus loko sp. nov.; Charinus longipes Weygoldt, 2006, stat. nov.; Charinus longitarsus Armas & Palomino-Cárdenas, 2016; Charinus madagascariensis Fage, 1954; Charinus magalhaesi sp. nov.; Charinus magua Seiter, Schramm & Schwaha, 2018; Charinus martinicensis Teruel & Coulis, 2017; Charinus milloti Fage, 1939; Charinus miskito sp. nov.; Charinus mocoa sp. nov.; Charinus monasticus sp. nov.; Charinus montanus Weygoldt, 1972; Charinus muchmorei Armas & Teruel, 1997; Charinus mysticus Giupponi & Kury, 2002; Charinus neocaledonicus Kraepelin, 1895; Charinus orientalis Giupponi & Miranda (2016); Charinus palikur sp. nov.; Charinus papuanus Weygoldt, 2006; Charinus pardillalensis (González-Sponga, 1998); Charinus pecki Weygoldt, 2006; Charinus perezassoi Armas, 2010; Charinus perquerens sp. nov.; Charinus pescotti Dunn, 1949; Charinus platnicki (Quintero, 1986); Charinus potiguar Vasconcelos, Giupponi & Ferreira, 2013; Charinus puri sp. nov.; Charinus quinteroi Weygoldt, 2002; Charinus reddelli Miranda, Giupponi & Wizen, 2016; Charinus renneri sp. nov.; Charinus ricardoi Giupponi & Miranda, 2016; Charinus rocamadre Torres-Contreras, García & Armas, 2015; Charinus ruschii Miranda et al., 2016; Charinus santanensis Vasconcelos & Ferreira, 2017; Charinus schirchii (Mello-Leitão, 1931); Charinus sillami Réveillion & Maquart, 2015; Charinus sooretama sp. nov.; Charinus souzai sp. nov.; Charinus spelaeus Vasconcelos & Ferreira, 2017; Charinus susuwa sp. nov.; Charinus taboa Vasconcelos, Giupponi & Ferreira, 2016; Charinus tingomaria Ballón-Estacio & Armas, 2019; Charinus tomasmicheli Armas, 2006; Charinus troglobius Baptista & Giupponi, 2002; Charinus tronchonii (Ravelo, 1975); Charinus una sp. nov.; Charinus vulgaris Miranda & Giupponi, 2011; Charinus wanlessi (Quintero, 1983) .

Distribution

Recorded from the following countries and territories: Australia, Belize, Brazil, Colombia, Cuba, Dominican Republic, Ecuador, Equatorial Guinea, French Guiana, Ghana, Guadeloupe, Guinea, Guiana, Jamaica, Kenya, Madagascar, Martinique, New Caledonia, Panama, Papua New Guinea, Puerto Rico, Saint-Barthélemy, Samoa, São Tomé and Príncipe, Sierra Leone, Surinam, US Virgin Islands, Venezuela.

Remarks

Twelve species previously assigned to Charinus are hereby transferred to Sarax based on the phylogeny of Miranda et al. (2021). Those species share with other species of Sarax the position of the lateral eyes situated near the carapace margin, the finger-like and plunger-like shape of the female gonopod, and the similar shape of the posterior margin of the genital operculum with denticulated surface on the dorsal area and ventral projections aligned with the gonopods.

Key to the identification of the species of Charinus in the Caribbean and Central America

1. Tibia of leg I with 23 articles............................................................................................................. 2

– Tibia of leg I with 21 or 22 articles.................................................................................................... 8

2. Leg I tarsus with 37–39 articles......................................................................................................... 3

– Leg I tarsus with 41 articles............................................................................................................... 4

3. Median eyes absent.................................................................. C. muchmorei Armas & Teruel, 1997

– Median eyes vestigial, reduced to small blackish spots; sternum platelets divided (Fig. 20E)........... ................................................................................................ C. cubensis (Quintero, 1983) (Fig. 20)

– Median eyes vestigial, reduced to small blackish spots; sternum platelets not divded ....................... .............................................................................................. C. martinicensis Teruel & Coulis, 2017

4. Median eyes present but reduced....................................................................................................... 5

– Median eyes absent............................................................................................................................ 6

5. Pedipalp femur with three dorsal spines................................................... C. acosta (Quintero, 1983)

– Pedipalp femur with two dorsal spines........................................ C. desirade Teruel & Questel, 2015

6. Lateral eyes reduced ............................................................ C. centralis Armas & Ávila Calvo, 2000

– Lateral eyes well developed............................................................................................................... 7

7. Leg I tarsus, first article 1.4–1.9 times as long as second article......................................................... ................................................................................... C. reddelli Miranda, Giupponi & Wizen, 2016

– Leg I tarsus, first article 3 or 4 times as long as second article ........................................................... ................................................................................ C. belizensis Miranda, Giupponi & Wizen, 2016

8. Leg I tarsus with 33 or 34 articles ..................................................................................................... 9

– Leg I tarsus with 37–39 articles........................................................................................................11

9. Median eyes present but reduced (Fig. 13A)............. C. aguayoi Moyá-Guzmán, 2009 (Figs 13–16)

– Median eyes absent.......................................................................................................................... 10

10. Pedipalp patella with three ventral spines (Fig. 21F) .......................................................................... .......................................................................... C. dominicanus Armas & Pérez, 2001 (Figs 21–23)

– Pedipalp patella with two ventral spines ............................................... C. wanlessi (Quintero, 1983)

11. Median eyes present ........................................................................................................................ 12

– Median eyes absent.......................................................................................................................... 13

12. Median eyes well developed; pedipalp femur with four dorsal spines................................................ ....................................................................................................... C. bruneti Teruel & Questel, 2011

– Median eyes reduced; pedipalp femur with three dorsal spines .......................................................... ................................................................................................ C. miskito sp. nov. (Figs 5E–F, 24–26)

13. Leg IV basitibia with two pseudo-articles..................... C. caribensis (Quintero, 1986) (Figs 17–19)

– Leg IV basitibia with three pseudo-articles ..................................................................................... 14

14. Leg I tarsus, first article three to five times as long as second article.............................................. 15

– Leg I tarsus, first article shorter or longer than three times second article ...................................... 16

15. Cheliceral claw with four teeth................................... C. magua Seiter, Schramm & Schwaha, 2018

– Cheliceral claw with five teeth ................................................................. C. perezassoi Armas, 2010

16. Tritosternum barely reaching base of pedipalp coxae ................................ C. bahoruco Teruel, 2016

– Tritosternum surpassing base of pedipalp coxae ............................................................................. 17

17. Leg I tarsus, first article 4.5 times as long as second article; leg I tarsus with 39 articles .................. ..................................................................................................................... C. decu (Quintero, 1983)

– Leg I tarsus, first article equal in length to second article; leg I tarsus with 37 articles...................... ............................................................................................................. C. tomasmicheli Armas, 2006