Systematic revision of the pantropical whip spider family Charinidae Quintero, 1986 (Arachnida, Amblypygi)
Author
Miranda, Gustavo Silva de
81150D94-592A-4CE5-8E88-E60F557A4341
Center for Macroecology, Evolution and Climate, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark. & Entomology Department, National Museum of Natural History, Smithsonian Institution, Washington DC, USA.
smiranda.gustavo@gmail.com
Author
Giupponi, Alessandro P. L.
434112AC-B212-43E8-A5D9-2F5D5619AFC4
Laboratório de Referência Nacional em Vetores das Riquetsioses, LIRN-IOC-FIOCRUZ, Rio de Janeiro, RJ, Brazil.
agiupponi@gmail.com
Author
Prendini, Lorenzo
C2D080D0-75DB-4DA1-A101-AB4DCF50FF0A
Arachnology Lab, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY, USA.
lorenzo@amnh.org
Author
Scharff, Nikolaj
F84D2235-66D2-460C-820D-80024068759D
Center for Macroecology, Evolution and Climate, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark. & Entomology Department, National Museum of Natural History, Smithsonian Institution, Washington DC, USA. & Zoology Section, Natural History Museum of Denmark, University of Copenhagen, Denmark.
nscharff@snm.ku.dk
text
European Journal of Taxonomy
2021
2021-09-24
772
1
409
http://dx.doi.org/10.5852/ejt.2021.772.1505
journal article
4042
10.5852/ejt.2021.772.1505
b65bb5a9-bbe7-49a4-af44-4b4c03121288
2118-9773
5536410
9B82A32F-0A07-47E3-8684-FED7C8EBF1E9
Genus
Sarax
Simon, 1892
Figs 11C–D
,
103–149
;
Tables 8–9
Sarax
Simon, 1892: 43
, 48
(
type
species:
Sarax brachydactylus
Simon, 1892
, by original designation).
Charinides
Gravely, 1911: 35
(
type
species:
Charinides bengalensis
Gravely, 1911
, by monotypy).
Syn. nov.
Phrynichosarax
Gravely, 1915b: 437
(
type
species:
Phrynichosarax cochinensis
Gravely, 1915b
, by original designation (synonymized by
Weygoldt 2000a: 25
)).
Lindosiella
Kritscher, 1959: 454
(
type
species:
Lindosiella ioannitica
Kritscher, 1959
, by monotypy).
Syn. nov.
Sarax
–
Kraepelin 1895: 45
;
1899: 250
. —
Pocock 1900: 131
. —
Gravely 1915b: 441
. —
Mello-Leitão 1931: 55
. —
Werner 1935: 471
. —
Weygoldt 2000a: 25
. —
Harvey 2003: 7
. —
Rahmadi
et al.
2010: 3
. —
Rahmadi & Kojima 2010: 476
.
Charinides
–
Gravely 1915b: 442
. —
Mello-Leitão 1931: 53
. —
Werner 1935: 471
. —
Quintero 1983: 27
. —
Weygoldt 2000a: 24
.
Phrynicosarax
–
Mello-Leitão 1931: 52
. —
Werner 1935: 470
.
Diagnosis
Sarax
may be distinguished from the other two genera of
Charinidae
by the following combination of characters: lateral eyes usually close to lateral margin of carapace (
Figs 105C
,
112C
); seta situated lateral to lateral ocular triad; dorsal carina on coxa with setiferous tubercles situated close to each other; opisthosoma with (Southeast Asian and Oceanian species) or without (species of Africa, the Middle East and South Asia) ventral sacs and ventral sac covers; female gonopod finger-like (
Fig. 138
), plunger-like (
Fig. 106
) or flat (
Fig. 146
); distal margin of female genital operculum with pair of projections aligned with gonopods (
Fig. 121A–B
); area of genital operculum between gonopods and posterior margin with denticulate projections in dorsal view (as opposed to smooth in
Charinus
and
Weygoldtia
;
Fig. 115D
).
Etymology
Unspecified by
Simon (1892)
; from the Greek word for a long, flowing garment.
Description
CARAPACE. Dorsoventrally compressed, wider than long, slightly bent downwards, ventral to lateral eyes; narrow median sulcus extending from median ocular tubercle to fovea. Anterior margin rounded, with six to eight anterior setae. Frontal process large, triangular, not visible in dorsal view. Three pairs of shallow sulci laterally on carapace, first pair of sulci slightly posterior to lateral boss; no sulci reaching midline; deep, oval fovea posteromedially. Median eyes and median ocular tubercle well developed, reduced or absent, depending on the species; two setae on median ocular tubercle. Lateral eyes well developed, reduced or absent, depending on the species, pale, small seta posterior to each lateral ocular triad; lenses directed anteriorly and dorsally.
STERNUM. Tetra-segmented, all platelets markedly sclerotized. Tritosternum rounded posteriorly and projected anteriorly into large, blunt tubercle, surpassing base of pedipalp coxae, with two anterior, two median, and two posterior setae, and seta between two posterior pairs, with smaller setae posteriorly. Medial platelet (tetrasternum) with single convex platelet, with pair of large setae anteriorly, and several small setae posteriorly. Third platelet (pentasternum) forming single convex platelet, smaller than medial platelet, with two setae anteriorly and some setae posteriorly. Sternites separated from each other by length of third platelet. No anterior setae in membranous region and four setae posteriorly.
OPISTHOSOMA. Oblong, tegument with nearly indistinguishable punctation. Ventral sacs and ventral sac cover present or absent, depending on the species.
GENITALIA. Female genital operculum broadly sclerotized between gonopods (projections) and posteromedian margin. Female gonopods finger-like (narrow and tubular) or plunger-like (narrow and tubular with apical invagination) with wrinkled base; wide plaque of genital operculum posterior to gonopods. Male gonopods soft, as wide as long; median lobes short and narrow, not surpassing fistula, lamellar; wide, rounded apex, almost reaching apex of lateral lobe; inner surface of gonopod spinous or glabrous.
CHELICERAE. Cheliceral sulcus with four prolateral teeth; first (dorsal) tooth bifid, dorsal cusp larger than ventral cusp. Third tooth slightly narrower and shorter than second tooth. Fourth tooth one-third longer than third tooth. Prolateral surface with two parallel, transverse rows of around fifteen small setae, ventrally to dorsally. Two setae, prolateral and retrolateral, on dorsodistal border, near membranous region of claw. Retrolateral surface of claw with row of setae, basally to dorsally. Claw with three to twelve denticles.
PEDIPALPS. Coxae dorsally with rounded carina in dorsal view, with or without seta inside carina and two or three setae on anterior margin. Trochanter with large, spiniform ventral apophysis with blunt apex pointing anteriorly, bearing 11–16 large setae and several smaller setae, at posterior margin; two subequal spines, one medially in anterior row of setiferous tubercles, other at retrolateral border, dorsal to apophysis, slightly curved prolaterally. Femur with two distinct setiferous tubercles proximal to spine 1; primary series with two to four dorsal spines; three to five ventral spines; setiferous tubercle proximal to ventral spine 1, between spine 1 and proximal margin. Patella with three to five dorsal spines; setiferous tubercle distal to spine I, about one-third length of spine I; spine I with three large and several small setae in proximal third; spine II with two large and some small setae in proximal third; spine III with one seta in distal third, one medially, and one proximally; spine IV (when present) one-third length of spine III with setae medially; spine V (when present) half length of spine IV, with seta medially. Two to four ventral spines, decreasing in length; setiferous tubercle between spine I and distal margin. Tibia with two dorsal spines, proximal spine two-thirds length of distal spine; ventral spine in distal half of tibia, with prominent setiferous tubercle close to its base; long setae between spine and distal margin. Tarsus with one to three dorsal spines. Cleaning organ about half length of tarsus article; ventral row of cleaning brush with 25–34 setae. Claw with long, curved, acute apex.
LEGS. All legs setose. Femur length on leg I>III>II>IV; ventral corner of prolateral surface of leg II–IV femora projecting into distinct spiniform process. Tibia of leg I with 21–28 articles; distal articles with two small trichobothria, dorsally and prolaterally; one trichobothrium on second, third and fourth (distal to proximal) articles, near distal margin, all situated dorsally; trichobothria absent on other articles. Leg I tarsus (basitarsus + distitarsus) with 37–41 articles covered with large number of sensilla. Leg IV basitibia with two to four pseudo-article. Leg IV distitibia with three proximal and 13–15 distal trichobothria. Leg IV basitibia–distitibia length DT>BT1>BT4>BT3>BT2.
Included taxa
Sarax abbatei
(
Delle Cave, 1986
)
comb. nov.
;
Sarax batuensis
Roewer, 1962
,
new status
,
Sarax bengalensis
(
Gravely, 1911
)
comb. nov.
;
Sarax bilua
sp. nov.
;
Sarax bispinosus
(
Nair, 1934
)
,
stat. nov.
;
Sarax brachydactylus
Simon, 1892
;
Sarax cavernicola
Rahmadi, Harvey & Kojima, 2010
;
Sarax cochinensis
(
Gravely, 1915
)
;
Sarax curioi
Giupponi & Miranda, 2012
;
Sarax dhofarensis
(Weygoldt, Pohl & Polak, 2002)
comb. nov.
;
Sarax dunni
sp. nov.
;
Sarax gravelyi
sp. nov.
;
Sarax huberi
Seiter, Wolff & Hörweg, 2015
;
Sarax indochinensis
sp. nov.
;
Sarax ioanniticus
(
Kritscher, 1959
)
comb. nov.
;
Sarax israelensis
(
Miranda
et al.
, 2016
) comb. nov.;
Sarax javensis
(
Gravely, 1915
)
;
Sarax lembeh
sp. nov.
;
Sarax mardua
Rahmadi, Harvey & Kojima, 2010
;
Sarax monodenticulatus
Rahmadi & Kojima, 2010
;
Sarax moultoni
(
Gravely, 1915
)
comb. nov.
, nomen dubium;
Sarax newbritainensis
Rahmadi & Kojima, 2010
;
Sarax omanensis
(
Delle Cave, Gardner & Weygoldt, 2009
)
comb. nov.
;
Sarax pakistanus
(
Weygoldt, 2005
)
comb. nov.
;
Sarax palau
sp. nov.
;
Sarax rahmadii
sp. nov.
;
Sarax rimosus
(
Simon, 1901
)
;
Sarax sangkulirangensis
Rahmadi, Harvey & Kojima, 2010
;
Sarax sarawakensis
(
Thorell, 1888
)
;
Sarax seychellarum
(
Kraepelin, 1898
)
comb. nov.
;
Sarax singaporae
Gravely, 1911
;
Sarax socotranus
(Weygoldt, Pohl & Polak, 2002)
comb. nov.
;
Sarax stygochthobius
(
Weygoldt & Van Damme, 2004
)
comb. nov.
;
Sarax timorensis
Miranda
& Reboleira, 2018;
Sarax tiomanensis
sp. nov.
;
Sarax willeyi
Gravely, 1915
;
Sarax yayukae
Rahmadi, Harvey & Kojima, 2010
,
Distribution
Cambodia
,
Greece
,
India
,
Indonesia
,
Israel
,
Italy
,
Jordan
,
Laos
,
Malaysia
,
Oman
,
Pakistan
,
Palau
,
Papua New Guinea
,
Philippines
,
Seychelles
,
Singapore
,
Solomon Islands
,
Somalia
,
Thailand
,
Timor-Leste
,
Turkey
,
Vietnam
,
Yemen
.
Remarks
Ten species previously assigned to
Charinus
are hereby transferred to
Sarax
based on the phylogeny of
Miranda
et al.
(2021)
. Those species share with other
Sarax
the position of the lateral eyes close to the carapace margin, the shape of the female gonopod (finger-like and plunger-like), and the similar shape of the posterior margin of the genital operculum with denticulate dorsal surface and ventral projections aligned with the gonopods.
Key to the identification of the species of
Sarax
in Southeast Asia and Oceania
1. Leg IV distitibia with six trichobothria in frontal and caudal series ................................................ 2
– Leg IV distitibia with five trichobothria in frontal and caudal series ...............................................11
2. Leg IV basitibia with three pseudo-articles ....................................................................................... 3
– Leg IV basitibia with four pseudo-articles ........................................................................................ 4
3. Leg IV distitibia, trichobothrium
bc
situated closer to
sbf
than to
bf
; pedipalp femur with setiferous tubercle between ventral spine 1 and proximal margin .........................
S. lembeh
sp. nov.
(
Fig. 117
)
– Leg IV distitibia, trichobothrium
bc
situated closer to
bf
than to
sbf
; pedipalp femur with spine between ventral spine 1 and proximal margin.......................................
S. singaporae
Gravely, 1911
4. Median and lateral eyes reduced........................................................................................................ 5
– Median and lateral eyes unmodified.................................................................................................. 7
5. Two pairs of lateral eyes ....................................................
S. timorensis
Miranda
& Reboleira, 2018
– Three pairs of lateral eyes.................................................................................................................. 6
6. Pedipalp femur with six dorsal spines and seven ventral spines ......................................................... ..............................................................................
S. cavernicola
Rahmadi, Harvey & Kojima, 2010
– Pedipalp femur with four dorsal spines and four ventral spines.......................................................... .....................................................................................
S. mardua
Rahmadi Harvey & Kojima, 2010
7. Pedipalp tarsus with one spine........................................
S. javensis
(
Gravely, 1915
)
(
Figs 115–116
)
– Pedipalp tarsus with three spines (
Fig. 131D
)..................................................................................... ..........................................................
S. yayukae
Rahmadi, Harvey & Kojima, 2010
(
Figs 131–134
) – Pedipalp tarsus with two spines......................................................................................................... 8
8. Carapace anterior margin anteriorly projected ...........
S. newbritainensis
Rahmadi & Kojima, 2010
– Carapace anterior margin rounded..................................................................................................... 9
9. Cheliceral claw with three teeth; cheliceral basal segment, retrolateral margin without tooth ........... .....................................................................................................
S. gravelyi
sp. nov.
(
Figs 110–111
)
– Cheliceral claw with five teeth; cheliceral basal segment, retrolateral margin with large tooth .... 10
10. Pedipalp patella with five or six dorsal spines...........................
S. rhamadii
sp. nov.
(
Figs 120–121
)
– Pedipalp patella with four dorsal spines ...
S. sangkulirangensis
Rahmadi, Harvey & Kojima, 2010
11. Cheliceral claw without teeth ...................................................
S. curioi
Giupponi & Miranda, 2012
– Cheliceral claw with three to five teeth ........................................................................................... 12
12. Carapace anterior margin anteriorly projected ................................................................................ 13
– Carapace anterior margin rounded................................................................................................... 14
13. Pedipalp patella with five dorsal spines (
Fig. 122E
); leg IV basitibia with three pseudoarticles...............................................................................
S. rimosus
(
Simon, 1901
)
(
Figs 122–124
)
– Pedipalp patella with three dorsal spines (
Fig. 130E
); leg IV basitibia with four pseudoarticles...........................................................................................
S. willeyi
Gravely, 1915
(
Fig. 130
)
14. Cheliceral basal segment, retrolateral margin without tooth .......................................................... 15
– Cheliceral basal segment, retrolateral margin with tooth ................................................................ 16
15. Pedipalp femur with three dorsal spines (
Fig. 125D
); pedipalp patella with four dorsal spines (
Fig. 125D
) .....................................................................
S. sarawakensis
(
Thorell, 1888
)
(
Fig. 125
)
– Pedipalp femur with four or five dorsal spines; pedipalp patella with three dorsal spines ................. ..........................................................................................................
S. bilua
sp. nov.
(
Figs 105–107
)
16. Cheliceral claw with two teeth ................................
S. monodenticulatus
Rahmadi & Kojima, 2010
– Cheliceral claw with three teeth ...................................................................................................... 17
– Cheliceral claw with four teeth........................................................................................................ 18
– Cheliceral claw with five teeth ........................................................................................................ 20
17. Female genital operculum concave with pair of bulges .......
S. tiomanensis
sp. nov.
(
Figs 126–129
)
– Female genital operculum flat ..........................................
S. brachydactylus
Simon, 1892
(
Fig. 108
)
18. Pedipalp patella with three dorsal spines.........................................
S. palau
sp. nov.
(
Figs 118–119
)
– Pedipalp patella with five dorsal spines........................................................................................... 19
19. Pedipalp tarsus with one spine.................................................................
S. dunni
sp. nov.
(
Fig. 109
)
– Pedipalpl tarsus with two spines..........................................
S. huberi
Seiter, Wolff & Horweg, 2015
20. Leg IV distitibia, trichobothrium situated
bc
closer to
bf
than to sbf ................................................. ...............................................................................................
S. batuensis
Roewer, 1962
(
Fig. 104
)
– Leg IV distitibia, trichobothrium situated
bc
midway between
bf
and
sbf ......................................... ..........................................................................................
S. indochinensis
sp. nov.
(
Figs 112–114
)