Thaumastella aradoides Horváth, 1896

Kment, Petr, Jacobs, Dawid H., Carapezza, Attilio, Deckert, Jürgen, Rider, David A. & Kóbor, Péter, 2024, Desert Bugs (Hemiptera: Heteroptera: Thaumastellidae): New records and review of the distribution and habitat of this relict group, Zootaxa 5541 (2), pp. 144-162 : 146-148

Thaumastella aradoides Horváth, 1896

( Figs 1–2 , 9, 11–12 )

Thaumastella aradoides Horváth, 1896: 325 (original description). Holotype: ♀, Algeria, Biskra (HNHM—see Péricart 1996: 153; Figs 1–2 ).

Thaumastella aradoides : Puton (1899): 28 (catalogue); Oshanin (1906): 289 (catalogue, systematic position); Oshanin (1912): 31 (catalogue); Stichel (1959): 324 (catalogue); Stichel (1960): 161 (catalogue); Seidenstücker (1960): 1 –4, Figs 1 –14 (redescription, morphology, systematic placement, collection circumstances, record); Seidenstücker (1964): 273 –278, Figs 9 –20 (morphology, systematic placement, collection circumstances, records); Slater (1964): 1498 (catalogue); Štys (1964): 239 –248, Figs 1–13 (morphology, systematic placement); Cobben (1968): 78 –80, Figs 66″A–C (description of egg); Schaefer (1975): 230, 235 (trichobothria); Linnavuori (1977): 49 (collection circumstances, records); Dolling (1981): 66, 70–71, Figs 16–20 (morphology, systematic placement, record); Schaefer (1981): 232 (morphology); Afzal & Sahibzada (1988): 254 (morphology); Linnavuori (1989): 6 (collection circumstances, record); Zrzavý (1990): 207, 208, 216, 219, 220, [222], Fig. 3e (morphology); Gapud (1991): 872, 878–879, 894 (morphology); Linnavuori (1993a): 142 –143, Figs 104p, 105a–h (figures, records); Linnavuori (1993b): 39 (record); Péricart (1996): 153 (type material); Hoberlandt (1997): 182 (collection circumstances, records) [paper omitted in Iranian catalogue by Ghahari et al. (2014)]; Linnavuori & van Harten (2002): 181 (record); Derjanschi & Péricart (2005): Pl. IV: Fig. D (habitus drawing); Linnavuori & van Harten (2006): 171 (records); Lis (2006): 117 (Palaearctic catalogue); Pluot-Sigwalt & Lis (2008): 298, 299, 306, 309, Fig. 66 (morphology of spermatheca); Kment & Vilímová (2010a): 6 (record); Kment & Vilímová (2010b): 5, 11, 12, 23, figs 12, 33–34, 101–102 (morphology of metathoracic scent gland efferent system, record); Ghahari et al. (2014): 84 (catalogue: Iran); Lis et al. (2017): 482, 484, Figs 1I, 2G (morphology, systematic placement, habitus photo); Rider et al. (2018): 135, 185, Fig. 2.16.J (systematic placement, habitus photo, record); Lis et al. (2024): 4, Fig. 1K , 5 , 6 , Fig. 2 (morphology, systematic placement, molecular phylogeny).

Type material examined. Holotype: ♀ ( Fig. 1 ), ALGERIA: ‘ Biskra [printed] // Thaumastella / aradoides Horv’ [handwritten by G. Horváth] ( HNHM).

Additional material examined. ASIA: OMAN: Al Batinah, 10 km W of Shinas , 24°44′N 56°23′E, 60 m a.s.l., 15.–16.iii.2017, 2 ♂♂ 2 ♀♀, S. Jakl lgt., A. Carapezza det. ( ACPI) .— SAUDI ARABIA: Jizan Province: 10 km SE of Al Eidabi, Wadi Eidabi , 17°09.657′N 42°59.520′E, 139 m a.s.l., 17.iii.2014, 47 ♂♂ 51 ♀♀, A. Carapezza lgt. & det. ( Fig. 2 , ACPI) .— UNITED ARAB EMIRATES: Sharjah Desert Park (25°17′N 55°42′E), light trap ([Sample No.] 12374), 2.iii.–1.iv.2007, 1 ♀, A. van Harten lgt., P. Kment det. ( NMPC) ; Wadi Bih (dam) (25°48′N 56°04′E), water trap (12962), 13.xii.2009 – 8.iii.2010, 2 ♂♂, A. van Harten lgt., P. Kment det. ( NMPC) ; Wadi Safad (25°13′N 56°19′E), light trap (12419), 22.iv.–21.vi.2006, 1 ♂, A. van Harten lgt., P. Kment det. ( NMPC) . YEMEN: Abyan Province: Al Kowd, ix.1999, 1 ♀, van Harten & Haruri leg., A. Carapezza det. ( ACPI) .— AFRICA: EGYPT: Giza Governorate: West Desert Oasis, Bahariya (28°21′5.36″N 28°51′44.6″E), light trap, 30.iv.1996, 1 ♀, W. G. Ulrich lgt., J. Deckert det. ( ZMHB) . SUDAN: Al Jazirah: Meddani [= Wad Madani] , Blue Nile, at light, 21.iii.1925, 1 ♀, H. W. Bedford lgt., P. Kment det. ( BMNH) . Kassala to Red Sea: Kassala Prov., Kassala – Haiya , 1.–3.xii.1962, 3 ♀♀, R. Linnavuori lgt. & det., P. Kment revid. ( BMNH) . Khartoum: Khartoum, 26.x.1965, 1 ♂, Omer lgt., P. Kment det. ( MMBC) ; Khartoum ([ Sample No. ] 28), [15.36°N 32.32°E, University, light trap], 14.ix.1965, 2 ♂♂ 3 ♀♀, P. Štys [on label, S. M. Omer in field diary] lgt. ( MMBC) ; Khartoum (47), [ University , light trap], 2.x.1965, 1 ♀, P. Štys [S. M. Omer] lgt., P. Kment det. ( MMBC) ; Khartoum (63), [University, light trap], 26.x.1965, 1 ♂ 2 ♀♀, P. Štys [S. M. Omer] lgt., P. Štys det., P. Kment revid. (1 ♂ 1 ♀ BMNH, 1 ♀ MMBC) ; Khartoum (463, 466, 467), [ University , light trap], 10.x.1966, 41 ♂♂ 115 ♀♀ 8 spec., P. Štys lgt., P. Kment det. (38 ♂♂ 106 ♀♀ 3 spec. MMBC; 4 ♂♂ 6 ♀♀> NMPC; 4 ♀♀> MNHN; 5 spec.> DARC [photographed in Rider et al. 2018: 185: Fig. 2.16J ]) [3 ♂♂ 22 ♀♀ 3 spec. in MMBC, incorrectly labelled as Khartoum-Burri (466)] ; Khartoum (479, 480), [ University , light trap], 21.x.1966, 11 ♂♂ 14 ♀♀, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum (1071), 7 [correctly 3]. x.1967, 1 ♀, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum (1072, 1073), [light trap], 3.x.1967, 2 ♂♂ 4 ♀♀, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum (1111), [ University , light trap], 31.x.1967, 2 ♂♂ 1 ♀, P. Štys lgt., P Kment det. ( MMBC) ; Khartoum-Burri (40), [15.36°N 32.34°E, Pink Palace Hotel, in bed], 20.–27.ix.1965, 1 ♂, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum-Burri (42), [little garden, at lamp], 27.–30.ix.1965, 2 ♀♀, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum-Burri (507), [little garden, at light], 2.xi.1966, 1 ♀, P. Štys lgt., P. Kment det. ( MMBC) ; Khartoum-Burri (1065), [at light], 1.–2.[correcty 3.] x.1967, 1 ♂, [P. Štys,] N. Williams & M. Hansell lgt., P. Kment det. ( MMBC) ; Shambat (375), [15.40°N 32.32°E, light trap in fields at the University Farm], 22.viii.1966, 1 ♂, P. Štys [on label, Venkatraman & Imam lgt. in field diary] lgt., P. Kment det. ( MMBC) ; Shambat (459), [light trap], 11.x.1966, 1 ♂, P. Štys lgt., P. Kment det. ( MMBC) ; Shambat (496), [light trap], 25.x.1966, 1 ♂, P. Štys lgt., P. Kment det. ( MMBC) . Sennar : Blue Nile Province, Dinder Game Reserve, Galegu (649), [12.36°N 35.02°E, at light], 27.ii.1967, 4 ♂♂ 8 ♀♀, P. Štys lgt., P. Kment det. ( MMBC) ; Dinder Game Reserve, Galegu (657), [at light], 28.ii.1967, 1 ♀, P. Štys lgt., P. Kment det. ( MMBC) . White Nile: Blue Nile Province, El Mellaha (631, 632, 633) [correctly White Nile, Ash Shawal , 13.32°N 32.38°E, light trap], 13.ii.1967, 5 ♂♂ 17 ♀♀, P. Štys lgt. & det., P. Kment revid. ( MMBC) ; Mal aba (637), [13.55°N 32.22°E, sweeping of ruderal vegetation + margin of field with hyacinth bean Lablab purpureus subsp. purpureus ], 14.ii.1967, 1 ♀, P. Štys lgt., P. Kment det. ( MMBC) .

Habitat. Very little is known about the biology and habitat of the species. According to Seidenstücker (1964), the species is an Eremian faunal element which occurs in the hot season of the year, swarming in the night, being attracted to light along with cydnids of the genera Aethus Dallas, 1851 , and Geotomus Mulsant & Rey, 1866 . Most of the available specimens were collected by light traps ( Seidenstücker 1960, 1964; Linnavuori 1977, 1989; Hoberlandt 1997; this paper). Interestingly, the species often co-occurs in the light traps with Camptocera glaberrima (Walker, 1872) ( Rhyparochromidae ) ( Seidenstücker 1960, Štys 1964, this paper). Linnavuori (1977, 1989) mentioned it was collected in sandy habitats. Seidenstücker (1960) hypothesized that it lives in epipsammon, an idea shared by Štys (1964), however, we lack any observation to confirm it. Seidenstücker’s (1960) prediction of existence of brachypterous morph of Th. aradoides has also not been confirmed, despite the brachypterous morph being the only known form in the three South African species ( Schaefer & Wilcox 1971, Jacobs 1989, this paper).

Hoberlandt (1997) summarized the information on its occurrence in Iran as follows: ‘ 2 ♂♂ and 2 ♀♀ from the vicinity of Kahkom, 20 km SW of Hajiabad, were collected by sweeping. Specimens from all other localities were collected by light trap mainly in coastal sandy plain (nos 202, 208, 209), dense savanna with tree-like growth with predominant Prosopis spicigera [= P. cineraria ], Acacia arabica [= Vachellia nilotica subsp. tomentosa ], Acacia nubica [= Vachellia oerfota var. oerfota ], Tamarix aphylla , Salvadora persica , Ephedra ciliata , Calligonum comosum , Calotropis procera . Collecting places, nos. 215 and 323 represented typical steppe zones.’ In the United Arab Emirates, Th. aradoides was collected by light or water traps in desert (Sharjah Desert Park) as well as mountainous areas (farms in Wadi Bih and Wadi Safad) (van Harten 2008 and this paper). Both in Oman and Saudi Arabia the collected specimens were attracted to light together with Camptocera glaberrima (Walker, 1872) ( Heteroptera : Rhyparochromidae ); in Oman they were collected in a sandy coastal plain with halophytic vegetation, in Saudi Arabia in an arid sandy area with scanty vegetation ( Vachellia , Phragmites , Tamarix ) close to a small water course. According to field diaries of P. Štys, all but one of his Sudanese specimens were collected at light (including the one found in a bed); the only exception is one female originating from sample collected by sweeping of ruderal vegetation at margin of a field with hyacinth bean Lablab purpureus subsp. purpureus ( Fabaceae ) in Mal’aba (this paper).

Host plant association. There is no evidence of a host plant. In case of the cited plant associations, they refer to surrounding vegetation or cases when the insect was stationary on the plant, i.e., no feeding behavior was observed.

Phenology. Based on the available collecting dates, the adults of Th. aradoides may be found all around the year. However, there seems to be differences between the particular regions as the answer to the local climatic conditions. In the Nile basin ( South Sudan, Sudan and Eritrea) and Chad the specimens were collected in the following periods: 5.–20.i., 13.–28.ii, 26.–27.iii., 10.–12.v., 20.–22.v., 5.vi., 22.viii., 14.ix.–31.x., 11.xi.–3.xii.; interestingly, the large series of specimens collected in Khartoum and its vicinity by Pavel Štys between 14.ix. and 31.x. has its distinct peak on 10.x. ( Seidenstücker 1964; Linnavuori 1977, 1993a; this paper). In Yemen, the specimens were collected from July to October ( Linnavuori 1989, 1993a; Linnavuori & van Harten 2002, 2006; this paper), while in the nearby Jizan ( Saudi Arabia), it was sampled on 17.iii. (this paper). In the United Arab Emirates and Oman, the specimens were collected on 2.iii.–1.iv., 22.iv.–21.vi., and 13.xii. (this paper), while all the records from the adjacent areas of Iran come from May ( Seidenstücker 1960, 1964; Hoberlandt 1997) and those from Baghdad ( Iraq) from September ( Seidenstücker 1964, Linnavuori 1993b). There is probably more than one generation per year, but their number cannot be estimated based on the available data.

Distribution ( Figs 9, 11 ). PALAEARCTIC REGION: Algeria ( Horváth 1896); Iran: Hormozgan ( Hoberlandt 1997, Kment & Vilímová 2010a,b), Kerman ( Seidenstücker 1960, 1964; misinterpreted as Sistan & Baluchestan in Ghahari et al. 2014); Iraq ( Seidenstücker 1964, Linnavuori 1993b); Oman (new record); Saudi Arabia (new record); United Arab Emirates (new record); Yemen ( Linnavuori 1989, 1993a; Linnavuori & van Harten 2002, 2006). AFROTROPICAL REGION: Egypt (new record); Chad ( Linnavuori 1993a); Eritrea ( Linnavuori 1993a); South Sudan ( Linnavuori 1977); Sudan ( Dolling 1981; Rider et al. 2018, without exact localities): Al Jazirah ( Linnavuori 1977), Kassala ( Linnavuori 1977), Khartoum (new record), Red Sea ( Linnavuori 1977), River Nile ( Linnavuori 1977), Sennar (new record), White Nile ( Linnavuori 1977, this paper).— Štys (1964) reported 1 ♂ and 4 ♀♀ from HNHM labelled as Saïgon, but considered its provenience from Vietnam as improbable (see also Schaefer & Wilcox 1971).

Most of the known localities of Th. aradoides are concentrated in three distinct areas: i) the Nile basin in South Sudan, Sudan and Eritrea; ii) Yemen and the adjacent Jizan province of the Saudi Arabia, and iii) areas around the Strait of Hormuz in the UAE, Oman and Iran. There are four exceptions—isolated localities Biskra in southern outskirts of Atlas Mts. ( Algeria), Farcha near N’Djamena east of Chad Lake ( Chad), West Desert Oasis ( Egypt), and Baghdad on Tigris River ( Iraq). Although the known localities are situated in Eremian region, the species apparently avoids the central desert areas of the Sahara, Arabian Peninsula and eastern Iran ( Fig. 12 ).

Figures

FIGURES 1–2. Habitus of Thaumastella aradoides Horváth, 1896. 1—holotype ♀ (2.40 mm) and its labels; 2—♀ (2.25 mm) from Saudi Arabia: Wadi Eidabi. (Photos: 1—P. Kóbor, 2—A. Carapezza).

FIGURES 9–11. Distribution maps of Thaumastella species: 9—general view; 10—detailed distribution of Th. aradoides Horváth, 1896 in North Africa and Near East; 11—detailed distribution of Th. elizabethae Jacobs, 1989, Th. namaquensis Schaefer & Wilcox, 1971 and Thaumastella sp. nov. in Namibia and South Africa.

FIGURES 12–13. Comparison of distribution of Thaumastella aradoides Horváth, 1896 (12) and the South African species of Thaumastella (13), Th. elizabethae Jacobs, 1989, Th. namaquensis Schaefer & Wilcox, 1971 and Thaumastella sp. nov., compared to ecoregions according to Shorthouse (2010).

FIGURES 3–5. Habitus of South African Thaumastella species. 3—Th. elizabethae Jacobs, 1989, holotype ♂ and its labels; 4—Th. namaquensis Schaefer & Wilcox, 1971, holotype ♂ and its labels; 5—living specimen of Th. namaquensis, Namibia, Gellap, 9.ix.2008. (Photos: 3–4—D. H. Jacobs, 5—J. Deckert).

FIGURES 6–7. Habitat of Thaumastella namaquensis Schaefer & Wilcox, 1971, South Africa, Farm Deurdrift near Springbok. (Photos: D. H. Jacobs).