Tijdschrift voor Entomologie 160 (2017) 75–88
The occurrence of stalk-eyed flies (Diptera, Diopsidae) in the Arabian Peninsula, with a review of cluster formation in the Diopsidae Hans R. Feijen*, Ralph Martin & Cobi Feijen
Catalogue and distribution data are presented for the six Diopsidae species known to occur in the Arabian Peninsula: Sphyracephala beccarii, Chaetodiopsis meigenii, Diasemopsis aethiopica, Diopsis arabica, Diopsis mayae and Diopsis sp. (ichneumonea species group). The biogeographical aspects of their distribution are discussed. Records of Diopsis apicalis and Diopsis collaris are removed from the list for Arabia as these were based on misidentifications. Synonymies involving Diasemopsis aethiopica and Diasemopsis varians are discussed. Only one out of four specimens in the D. elegantula type series proved conspecific with D. aethiopica. The synonymy of D. aethiopica and D. varians is rejected. A lectotype for Diasemopsis elegantula is now designated. D. elegantula is proposed as junior synonym of D. varians. A fly cluster of more than 80,000 Sphyracephala beccarii, observed in Oman, is described. The occurrence of cluster formations in the Diopsidae is reviewed, while a possible explanation is indicated. Hans R. Feijen*, Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands. [email protected] Ralph Martin, University of Freiburg, Münchhofstraße 14, 79106 Freiburg, Germany Cobi Feijen, Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands
Introduction catalogue for Diopsidae, Steyskal (1972) only re- Westwood (1837b) described Diopsis arabica as ferred to Westwood and Hennig as far as Diopsidae the first stalk-eyed fly from the Arabian Peninsula. in Arabia was concerned. Feijen (1989) recorded As type locality only “in Arabiâ desertâ” was given. Sphyracephala beccarii (Rondani) and Chaetodiop Hennig (1941), in a contribution to “Die Fliegen sis meigenii (Westwood) for the Arabian Peninsula. der palaearktischen Region”, gave a partial redescrip- Feijen mentioned the occurrence of seven Diopsi- tion of D. arabica based on the type. Hennig indi- dae species for this peninsula, but one of these later cated that it was not certain whether the type was turned out to be based on a wrong geographic inter- collected in the Palaearctic or the Ethiopian part of pretation of a label. Arabia. He was anyway of the opinion that it did In the past ten years, five papers were published not concern a typical representative of the Palaearctic dealing with the Diopsidae fauna of Arabia. In the fauna, but, at most, a species which had penetrated first of these papers, Dawah & Abdullah (2008) re- from the Ethiopian Region. The type now appears to corded three Diopsidae species from south-western be lost (Rohlfien & Ewald 1970). In the last world Saudi Arabia: “Diopsis collaris”, “Diopsis apicalis” and
Tijdschrift voor Entomologie 160: 75–88, Figs 1–21. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 December 2017. DOI 10.1163/22119434-00002065 *Corresponding author Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Sphyracephala beccarii. In the second one, Hauser Diopsis beccarii Rondani, 1873: 289 – type series et al. (2011) discussed the occurrence of Sphyraceph from Eritrea, Sciotel, Bogos, 15°35’N 38°20’E, ala beccarii in the United Arab Emirates. They also lectotype and part of large type series in MCSN, gave records for the same species from Oman and paralectotypes in many other museums. Yemen. In the third paper, El-Hawagry et al. (2013) Hexechopsis beccarii, Rondani 1875: 442. Ref.: Osten listed “Diopsis apicalis” and Sphyracephala beccarii for Sacken 1882. Al-Baha Province, Saudi Arabia. In El-Hawagry et al. Sphyracephala beccarii, Osten Sacken 1882: 235. Ref.: (2015) “Diopsis collaris” was added to the list for Al- Steyskal 1972, Feijen 1989, Hauser et al. 2011. Baha Province, Saudi Arabia, while El-Hawagry et al. Diopsis africana Karsch, 1888: 380 – holotype from (2016) added more records for “Diopsis apicalis” and Tanzania, Bondei, ~5°00’S 39°00’E, in ZMHB Sphyracephala beccarii. but lost (see Rohlfien & Ewald, 1970). Ref.: Cur- We present here an annotated catalogue and a key ran 1928, Séguy 1949, Collart 1954, Steyskal to the six species presently recognized as occurring 1972, Cogan & Shillito 1980. in Arabia. As the border between the Palaearctic Re- gion and the Ethiopian Region in the Arabian Pen- Distribution. Algeria, all contiguous Sub-Saharan insula is formed by a somewhat indeterminate line, African countries, Madagascar, Arabian Peninsula. the biogeographical implication of the Diopsidae Arabian records. Oman, 7♀, 3♂, Wadi Siya, 23°12’N records will be discussed briefly. In Diopsidae, clus- 58°41’E, 400 m, 10.iv.1980, B.R. Pitkin (BMNH); tering was only known in the genus Sphyracephala. >80,000 specimens, Wadi Darbat, 17°6’58”N Feijen (1989) briefly reviewed the scarce records for 54°27’18”E, 206 m, 10.xi.2014, R. Martin, M. this phenomenon in stalk-eyed flies. A recent obser- Kühn, T. Wulf (observation & photographs); 2♀, vation of clustering in Oman is described. A review 1♂, Ain Razat, 17°7’44"N 54°14’11”E, 200 m, is presented for clustering in the Diopsidae while a 13.ii.1989, M.J. Ebejer (Hauser et al. 2011) possible explanation will be indicated. (NMWC); 2♀, Dhofar, Ain Sahwoot, 9.xi.1992, J.C. Deeming (Hauser et al. 2011) (NMWC); 2♀, Museum codens used 2♂, Dhofar, Ain Jarziz, 17°06’N 54°05’E, 197 m, 11.xi.1992, J.C. Deeming (Hauser et al. 2011) (NMWC); 1 Dhofar, Ain Hamran, 17°6’4”N BMNH The Natural History Museum [formerly British ♂, Museum (Natural History)], London, United 54°16’37”E, 1.xi.1988, M.J. Ebejer (Hauser et al. Kingdom 2011) (NMWC); 1♂, Dhofar, Hajayf, 17°17’N ISNB Institut Royal des Sciences Naturelles de 54°3’E, 12.x.1990, J.C. Deeming (Hauser et al. Belgique, Brussels, Belgium 2011) (NMWC); 1 , Zufar, near Ayn Razat, KSMA King Saud Museum of Arthropods, King Saud ♀ University, Riyadh, Saudi Arabia. 17°7’44”N 54°14’11”E, 200 m, 24.v.2012, Moayed MCSN Museo Civico di Storia Naturale Giacomo Bahajjaj (photographic record, Flickr); Ayn Razat, Doria, Genova, Italy 17°7’44”N 54°14’11”E, 200 m, 28.vi.2012, Moayed MNHNP Muséum National d’Histoire Naturelle, Paris, Bahajjaj (photographic record, Flickr); Ain Garziz, France MRAC Musee Royal de l’Afrique Centrale, Tervuren, Zufar, 17°6’N 54°5’E, 197 m, 7.iii.2012, Faisal Sa- Belgium lim (photographic record, Flickr); Saudi Arabia, 2♀, NHMB Naturhistorisches Museum, Basel, Switzerland 2♂, Wadi Maharish, 21°21.5’N 40°13’E, 1000 m, NHMW Naturhistorisches Museum Wien, Vienna, 13.i.1983, W. Büttiker (RMNH); 20♀, 28♂, Austria NHRS Naturhistoriska Riksmuseet, Stockholm, Maraba, Asir, 17°54’N 42°23’E, 80 m (or 350 m?), Sweden 2004–2005 (Dawah & Abdullah 2008) (NMWC, NMWC National Museum of Wales, Cardiff, United NHMCSA); Al-Mekhwa, 19°46’14”N 41°26’3”E, Kingdom 350 m, xii–ii.2008–2012 (El-Hawagry et al., 2013) NHMCSA Natural History Museum, College of Science at (KSMA); Dhee Ain, 19°33’16”N 41°15’46”E, Abha, Saudi Arabia. OXUM Hope Entomological Collections, Oxford 155 m, ii–v.2008–2012 (El-Hawagry et al., 2013) University Museum, Oxford, United Kingdom (KSMA); Jabal Shada al-A’la Nature Reserve, RMNH Naturalis Biodiversity Center (formerly 19°51’46”N 41°18’5”E 2.vi.2014, 19°50’20–43”N Rijksmuseum van Natuurlijke Historie), 41°18’16–41”E 3.vi.2014, 19°50’43”N 41°18’16”E Leiden, The Netherlands ZMHB Museum für Naturkunde der Humboldt- 27.i.2015, 19°50’35”N 41°18’42”E 5.v.2015, Universität, Berlin, Germany 19°51’46”N 41°18’5”E, 15.xi.2015 (El-Hawagry et al., 2016) (KSMA); Yemen, 12♀, 14♂, Wadi Dareija, SW of Dhala, W. Aden Prot., 13°40’45”N Annotated catalogue of Diopsidae in the 44°42’7”E, 4500’, 6–9.xi.1937, H. Scott & E.B. Arabian Peninsula Britton; 1♀, Al-Lahima, 15°24’N 43°32’E, 1200 m, Sphyracephala beccarii (Rondani) 16.x–31.xii.2000, A. van Harten & A.M. Hager Figs 1, 5, 11, 16–18, 20 (Hauser et al. 2011) (NMWC); 1♀, Suq Bani Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Figs 1–4. Distribution maps of Diopsidae species in the Arabian Peninsula. – 1, Sphyracephala beccarii (also show- ing type locality in Eritrea); 2, Chaetodiopsis meigenii and Diasemopsis aethiopica (also showing type locality in Eritrea); 3, Diopsis mayae; 4, Diopsis arabica and Diopsis sp.
Mansour, 15°6’N 43°50’E, 1500 m, 28.viii–26. are mapped in Fig. 1, which also shows the type ix.2001, A. van Harten (Hauser et al. 2011) locality. (NMWC); 1♀, Ar-Rujum, 15°27’28”N 43°38’6”E, 1989 m, 16.x.2000, A. van Harten & A.M. Hager Chaetodiopsis meigenii (Westwood) (Hauser et al. 2011) (NMWC); 1♀, 1♂, 12 km NW of Figs 2, 8, 12, 18–19 Manakhah, 15°6’N 43°43’E, 1500 m, 21.viii.2001, A. van Harten (Hauser et al. 2011) (NMWC); 1♀, Diopsis meigenii Westwood, 1837b: 548 (as Miege 12 km NW of Manakhah, 15°6’N 43°43’E, 1500 m, nii) – in Guineâ Africæ, type series (1♀, 1♂) in 5.v–17.vi.2002, A. van Harten (Hauser et al. 2011) OXUM. Ref.: Eggers 1925. (NMWC); United Arab Emirates, 21♀, 26♂, Wadi Diopsis subfasciata Macquart, 1843: 395 – errone- Hayl, 25°04.83’N 56°13.53’E, 240 m, 15.iii.2008, ously described from Java, Indonesia, holotype in M. Hauser, J-H. Stuke; 1♀, Wadi W Mirba, MNHNP (fragment). Ref.: Feijen 1978. 25°16.22’N 56°16.68’E, 260 m, 13.iii.2008, J.-H. Diopsis leucochira Bezzi, 1908a: 387 – type series Stuke; 1♂, Wadi Shawkah, 25°6’N, 56°2’E, 309 from DR Congo & South Africa, in ISNB. Ref.: m, 20–26.iii.2007, F. Menzel. Distribution records Feijen 1978, Feijen 1989. Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Figs 5–10. Anterior view of head or central head. – 5, Sphyracephala beccarii, female, Saudi-Arabia, Wadi- Maharish; 6, Diopsis mayae, female, Mozambique, Namaacha; 7, Diopsis mayae, male, South Africa, Pietermaritz- burg; 8, Chaetodiopsis meigenii, female, Namibia, Katima; 9, Diasemopsis aethiopica, male, DR Congo, Mboma; 10, Diopsis arabica, female, Saudi-Arabia, Jebel Fifa. Scale bars = 0.5 mm, photographs Cobi Feijen.
Not Diopsis breviseta Bezzi, 1908b:167 – Eritrea, Distribution. All contiguous Sub-Saharan African Massaua, Abd el Kader, 15°37’26”N 39°28’25”E. countries, Arabian Peninsula. Ref.: Lindner 1962, Feijen 1978, Cogan & Shil- Arabian records. Oman, Wadi Darbat, 17°6’58”N lito 1980, Feijen 1989, Carr et al. 2006. 54°27’18”E, 206 m, 10.xi.2014, R. Martin, M. Diasemopsis meigenii, Brunetti 1926a: 173. Ref.: Kühn & T. Wulf (observation & photographs); Cogan & Shillito 1980, Carr et al. 2006. 1♀, Kelen, Zufar, 17°5’16”N 54°19’10”E, 120 m, Chaetodiopsis meigenii, Séguy 1955: 1108. Ref.: 28.ix.2012, Moayed Bahajjaj (photographic record, Steyskal 1972, Feijen 1989. Flickr); Saudi-Arabia, 1♀, Fifa (Fayfa?), nr. Gizan, Downloaded from Brill.com10/06/2021 02:46:57PM via free access
17°15’N 43°6’E, 1240 m, 27–31.iii.1983, C. Holz D. veliventris and D. elegantula as synonyms of schuh (NHMB); Yemen, 16♀, 12♂, Usaifira, 1 mile D. aethiopica. The main illustration and the descrip- n. of Ta’izz, 13°38’N 44°1’E, 4500 ft, 13.xii.1937, tion by Eggers (1916) clearly cover D. varians as now H. Scott & E.B. Britton (BMNH); 1♂, Ta’izz, distinguished from D. aethiopica. Eggers considered 13°38’N 44°1’E, i.1951 (NHMB). Distribution re- D. varians as rather variable and related to D. aethi cords are mapped in Fig. 2. opica. From the description and the two additional abdomens illustrated, it is clear that Eggers’ type se- Diasemopsis aethiopica (Rondani) ries of six specimens must have been mixed, but it Figs 2, 9, 13 did not contain a D. aethiopica. Whether the type series still exists is not clear. Diopsis aethiopica Rondani, 1873: 289 – holotype Van Bruggen (1961) thought it convenient to ac- from Eritrea, Sciotel, Bogos, 15°35’N, 38°20’E, cept the redescription given by Curran (1931), but in MCSN Ref.: Guiglia 1957. did not wholly agree with his synonymy. He con- Diasemopsis aethiopica, Rondani 1875: 442. sidered D. elegantula Brunetti as certainly not syn- Diasemopsis veliventris Hendel, 1923: 41 – Kenya, onymous with Rondani’s species. Examination of the Wa-Kikuyu, Tchania (Chania) River and Wa- three “cotypes” of D. elegantula in MRAC revealed Taita, Bura, in NHMW. Ref.: Curran 1931, van that two are conspecific withD. varians while one Bruggen 1961, Steyskal 1972. is conspecific with D. aethiopica. The “syntype” in Not Diasemopsis varians Eggers, 1916: 12, 27 – Ke- BMNH is rather incomplete (no abdomen) and is nya, River Lumi (type series not traced). Ref.: likely to be also a D. varians. One of the two MRAC Curran 1931, van Bruggen 1961, Steyskal 1972. flies conspecific with D. varians is here designated Not Diasemopsis elegantula Brunetti, 1926b: 82 – type as the lectotype of D. elegantula, while the second series consisting of 3 “cotypes” from DR Congo specimen (no head) is designated as paralectotype. (MRAC) and 1 “syntype” from Durban, South Af- The third MRAC fly, conspecific withD. aethiopica, rica (BMNH). Three specimens of the type series, also becomes a paralectotype. D. elegantula is now including the now newly designated lectotype, proposed as new junior synonym of D. varians. are conspecific withD. varians, while only one paralectotype from DR Congo is conspecific with Diopsis mayae Feijen & Feijen D. aethiopica. Ref.: Curran 1931, van Bruggen Figs 3, 6–7, 14 1961, Steyskal 1972. Diopsis mayae Feijen & Feijen, 2017: 62 – holotype Distribution. All contiguous Sub-Saharan African from Malawi, Kasungu National Park, Lifupa, countries, Arabian Peninsula. 13°3’24”S 33°9’26”E in RMNH, large series of Arabian record. 1♀, Yemen, Wadi Tiban, NW of paratypes from Malawi, Mozambique and Tanza- Jebel Jihaf, 13°47’N 44°34’E, 3800 ft, 21.x.1937, nia in RMNH. H. Scott & E.B. Britton. Distribution record is Not Diopsis apicalis Dalman, 1817: 216 – holotype mapped in Fig. 2, which also shows the type locality. from Sierra Leone, in NHRS. Ref.: Feijen 1987. Remarks. Diasemopsis aethiopica and D. varians are Diopsis apicalis, Dawah & Abdullah 2008 two of the most common and abundant Diasemopsis (misidentification). with a wide distribution in the Afrotropical region. Diopsis apicalis, El-Hawagry et al. 2013 They often occur together. Although closely related, (misidentification). they can easily be separated. Diasemopsis varians is Diopsis apicalis, El-Hawagry et al. 2016 smaller, darker and a bit more hairy. The first ster- (misidentification). num is grey in D. aethiopica and black in D. vari ans. The pollinose spots on the abdomen are angu- Distribution. Angola, Botswana, Central African lar in D. varians and rounded in D. aethiopica. In Republic, DR Congo, Egypt, Ethiopia, Kenya, D. aethiopica the wing has a vague central wing spot Malawi, Mozambique, Namibia, Saudi Arabia, (Fig. 13), while in D. varians the wing is apically Somalia, South Africa, South Sudan, Swaziland, slightly infuscated. Being such common species, it Tanzania, Uganda, Yemen, Zambia, Zimbabwe. is not surprising that they have several times been Arabian records. Saudi Arabia, 1♀, 1♂, Wadi Min- described as new species. Curran (1931) considered sah, 20°41’N 40°40’E, 550 m, 7–8.iv.1983, W. Büt- D. aethiopica “A variable species in the female sex”. tiker (RMNH); 2♀, Asir, Jaheri, 16°33’N 43°3’E, He indicated a colour difference between males and 11.iii.1971, G. Popov (BMNH); 4♀, 1♂, Asir, females: “Male – abdomen rarely reddish on the Wadi Hali, 18°36’N 41°18’E, 420 m?, 9.i.2003, fourth segment, otherwise like the female”. However, H.A. Dawah (Dawah & Abdullah 2008) (NMWC, reddish-brown fourth sterna and especially third ster- NHMCSA); 1♀, 1♂, Asir, Maraba, 17°54’N 42°23’ na occur in both sexes. Curran indicated D. varians, E, 350 m? (Dawah & Abdullah 2008) (NMWC, Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Figs 11–15. Dorsal view of wing. – 11, Sphyracephala beccarii, female, Saudi-Arabia, Wadi-Maharish; 12, Chae todiopsis meigenii, male, DR Congo, Bomane; 13, Diasemopsis aethiopica, female, Togo, Amoutchou; 14, Diopsis mayae, male, Mozambique, Buzi; 15, Diopsis arabica, female, Saudi-Arabia, Jebel Fifa. Scale bars = 1 mm, photo- graphs Cobi Feijen.
NHMCSA); 5♀, 4♂, Albahha, De Al-Ain, 20°0’N (KSMA); Yemen, 5♀, 1♂, Mukeras, (Mukayris?), 41°28’E, 3.iv.2003 (Dawah & Abdullah 2008) 13°56’N 45°40’E, 2159 m, i.1936, R.C.M. Dar- (NMWC, NHMCSA); Al-Mekhwa, 19°46’14”N ling (BMNH); 2♀, Wadi Lejij, Jebel Jihaf, 13°44’N 41°26’3”E, 350 m, xii–ii.2008–2012 (El-Hawagry 44°40’E, 7000 ft, 1.x.1937, H. Scott & E.B. Britton; et al., 2013) (KSMA); Dhee Ain, 19°33’16”N 3♀, 2♂, Wadi Dareija, SW of Dhala, 13°40’45”N 41°15’46”E, 155 m, ii–v.2008–2012 (El-Hawagry 44°42’7”E, 4500 ft, 6/9.ix.1937, H. Scott & E.B. et al., 2013) (KSMA); Jabal Shada al-A’la Nature Britton (BMNH); 9♀, 6♂, Usaifira, 1 mile N of Reserve, 19°50’35”N 41°18’42”E and 19°50’25”N Ta’izz, 13°38’N 44°1’E, 4500 ft, 21.xii.1937, 41°18’41”E, 5.v.2015 (El-Hawagry et al., 2016) H. Scott & E.B. Britton (BMNH); 8♀, 8♂, 1?, Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Figs 16–19. Wadi Darbat, Sultanate of Oman, 10.xi.2014. – 16, habitat of the Wadi; 17, overview of clusters of Sphyracephala beccarii on tree trunk, dark sections are clusters, specks on pale sections are individual flies; 18, detail of cluster of Sphyracephala beccarii, single Chaetodiopsis meigenii in lower left corner; 19, Chaetodiopsis meigenii near cluster. Photographs 16, 17 Martin Kühn; photographs 18, 19 Ralph Martin.
Wadi Jaira, tributary of Wadi Siham, 14°52’N Diopsis arabica Westwood, 1837b: 544 – in Arabiâ 43°46’E, 3000 ft, 10.iii.1938, H. Scott & E.B. Brit- desertâ. Type in ZMHB lost after 1941 (see Rohl- ton (BMNH). Distribution records are mapped in fien & Ewald 1970). Ref.: Hennig 1941, Steyskal Fig. 3. 1972, Cogan & Shillito 1980, Feijen 1989. Remarks. As indicated by Feijen (1987) and Feijen & Teleopsis arabica, Rondani 1875: 443. Ref.: Brunetti Feijen (2017), many of the recordings for D. apica 1928, Hennig 1941, Shillito 1971. lis (or its junior synonym D. tenuipes Westwood) are Diopsis collaris, Dawah & Abdullah 2008 misidentifications, involving one of the few other de- (misidentification). scribed Diopsis with an apical wing spot, or one of the Diopsis collaris, El-Hawagry et al. 2015 more than 60 undescribed Diopsis with such a wing (misidentification). spot. Diopsis apicalis, as defined by Feijen 1987 and Fei- jen & Feijen (2017), has a West-African distribution Distribution. Arabian Peninsula, but perhaps also from Mauritania to Chad and to Gabon. It is the domi- Eritrea. nant West-African species of the D. apicalis species Arabian records. Saudi Arabia, 1♀, 1♂, Asir, Jaheri, group (Feijen & Feijen 2012). In eastern and southern 16°33’N 43°3’E, 11.iii.1971, G. Popov (BMNH); Africa it is replaced by the also very abundant D. mayae, 1♀, Fifa, nr. Gizan, Jebel Fifa, 17°15’N 43°06’E, which even expands into Egypt and the Arabian Penin- ~1240 m, 15.vii.1981, A.S. Talhouk (NHMB); 2♂, sula. Diopsis mayae is the sister species of D. apicalis. Asir, Wadi Hali,18°36’N 41°18’E, 420 m? 9.i.2003 (Dawah & Abdullah 2008) (NMWC, NHMCSA); Diopsis arabica Westwood 3♀, 2♂, Albahha, De Al-Ain, 20°0’N 41°28’E, Figs 4, 10, 15 3.iv.2003 (Dawah & Abdullah 2008) (NMWC, Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Figs 20–21. Clusters in Diopsidae. – 20, Wadi Darbat, Sultanate of Oman, 10.xi.2014, detail of cluster of Sphy racephala beccarii showing layered structure (photograph Ralph Martin); 21, Matema, Tanzania, 6.vi.2014, cluster of Diasemopsis varians with some Sphyracephala munroi in between, note densely layered structure on lower leaf just left of centre (photograph Martin Grimm).
NHMCSA); 13♀, 7♂, Maraba, Asir, 17°54’N referred to as the D. ichneumonea species group 42°23’E, 15.viii.2004, 15.x.2004, 17.xi.2004, (Feijen & Feijen 2009) and counts 13 recognized 15.xii.2004, 4.ii.2005, 17.iii.2005, 3.v.2005 (Dawah African species and D. arabica. Most species in this & Abdullah 2008), (NMWC & NHMCSA); group remain to be described. Four of the presently Yemen, 6♀, 2?, Usaifira, 1 mile N of Ta’izz, 13°38’N recognized species have a red collar: D. ichneumonea 44°1’E, 4500 ft, 21.xii.1937, H. Scott & E.B. Brit- Linnaeus, D. pallida Westwood, D. collaris West- ton (BMNH); 3♀, 1♂, Wadi Jaira, tributary of Wadi wood and D. arabica. Important external characters Sihan, 14°52’N 43°46’E, 3000 ft, 10.iii.1938, H. in this group are: shape of body (slender forest spe- Scott & E.B. Britton (BMNH); 1♀, 1♂, 1?, Wadi cies or more broadly built species from open coun- Tiban, NW of Jebel Jihaf, 13°47’N 44°34’E, 3800 try), shape of preapical wing spot, pollinosity pat- ft, 21.x.1937, H. Scott & E.B. Britton (BMNH). tern of thorax, external structure of scutum (smooth Distribution records are mapped in Fig. 4. or granular) and colour of collar. As far as we know Remarks. In Africa the number of Diopsis species there are no species in this group with a large distri- with a preapical wing spot is high. This group is bution. West African species do not occur in East Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Africa and vice versa. Westwood (1837a) described – Front femur only with two rows of tubercles; D. collaris from Senegal while his D. arabica came apical seta absent on scutellar spine; abdo- from Arabia. The thorax of D. collaris he described men yellowish to reddish brown without as largely “niger, nitidus”, while the thorax of D. ara pollinosity patterns; three spermathecae; bica was described as “sericeo-niger”, i.e. pollinose. DIOPSIS ���������������������������������������������������� 4 Feijen (1989) indicated that the “D. ichneumonea” 3. Central head almost rectangular in anterior from Eritrea recorded by Frey (1921) could have view, face black and smooth, distinct facial been a D. arabica. The Arabian species identified by teeth, frons with ribbed structure centrally Dawah & Abdullah (2008) and El-Hawagry et al. (Fig. 8); wing with narrow central band (in- (2015) as D. collaris should now be regarded as re- cluding two darker spots) and small apical cords for D. arabica. spot (Fig. 12); abdomen distally broad and rounded; distinct suture between surstyli and Diopsis sp. epandrium ���������������Chaetodiopsis meigenii Fig. 4 – Central head distinctly tapering ventrally (al- most triangular in males with large stalks), Diopsis sp. This species also belongs to the D. ich frons and face brownish with paler brown neumonea species group, but it could not be as- spots, no facial teeth, frons smooth centrally signed to a described species and is most likely un- (Fig. 9); wing with very vague central and api- described. However, describing and redescribing cal infuscation (Fig. 13); abdomen slender, ta- the many species (>50) in this probably not even pering apically; surstyli seamlessly fused with monophyletic species group is a major job. epandrium �������������� Diasemopsis aethiopica 4. Central head trapezoid (Fig. 6); wing with Distribution. Possibly Arabian Peninsula only. large rounded apical wing spot (Fig. 14); scu- Arabian records. Yemen, 1♀, 2♂, Usaifira, 1 mile N tum largely glossy; surstyli fused to epandrium of Ta’izz, 13°38’N 44°1’E, 4500 ft, 21.xii.1937, H. �����������������������������������������������Diopsis mayae Scott & E.B. Britton (BMNH); 3♀, 1♂, Jebel Jihaf, – Central head rounded (Fig. 10); wing with Wadi Leje, 13°27’N 44°26’E, 6800 ft, 13–15.x.1937, preapical wing spot (Fig. 15); scutum largely H. Scott & E.B. Britton (BMNH); 1♀, Wadi Jaira, thinly pollinose; surstyli articulate ���������������� 5 tributary of Wadi Sihan, 14°52’N 43°46’E, 3000 ft, 5. Collar reddish brown �����������Diopsis arabica 10.iii.1938, H. Scott & E.B. Britton (BMNH). Dis- – Collar black �������������������������������������������������� tribution records are mapped in Fig. 4. �������� Diopsis sp. (ichneumonea species group) Remarks. This Diopsis from the ichneumonea species group can readily be distinguished from D. arabica Biogeographical aspects of the Diopsidae in by its black collar. In D. arabica the collar is reddish Arabia brown. Colour of the collar is a major character in Diopsidae chiefly occur in the Afrotropical and this species group. Oriental Regions. In the Nearctic Region two spe- cies of Sphyracephala occur (Feijen 1989). In the Australasian Region undescribed Sphyracephala A key to the Diopsidae in Arabia species occur in the Solomon Islands and on New 1. Arista tripartite; eye stalk stout (~0.70× the Guinea. On Taiwan, with both Palaearctic and widest sagittal eye diameter, Fig. 5), alula pres- Oriental aspects, two species occur: Megalabops ent (Fig. 11); vein CuA+CuP extending past bigotii (Hendel) and an undescribed Sphyracephala cell cua; apical seta more than three times the from the detrahens species group (see Feijen 1989). scutellar spine length; syntergum only with Likewise in the biogeographically hybrid Yaeyama terga 1+2 �����������������Sphyracephala beccarii Islands of Japan an undescribed Sphyracephala from – Arista bipartite; eye stalk slender (<0.30× the the detrahens species group occurs (Ôhara 1993). In eye diameter, Fig. 7), alula absent (Figs 12–15); the undisputed Palaearctic Region three Sphyraceph vein CuA+CuP not extending past cell cua; ala species occur in, respectively, 1) Hungary and apical seta usually shorter than scutellar spine Serbia, 2) Azerbaijan and 3) Russia (Primorsky Krai) or absent; syntergum including terga 1+2+3; and Manchuria. A record of an unidentifiedDiopsis DIOPSINAE ���������������������������������������������� 2 in Greece (Anagnou-Veroniki et al. 2008), probably 2. Front femur with two rows of spinous se- represents an isolated case of introduction into the tae around two rows of tubercles; apical seta rice fields. From Europe, extinct Prosphyracephala are present on scutellar spine; abdomen dark known from both amber and oil-shale sediments (see with pattern of silvery-grey pollinosity; two Kotrba 2009). spermathecae; DIASEMOPSIS GENUS From the Afrotropical Region two intrusions of GROUP ������������������������������������������������������ 3 Diopsidae into Palaearctic North Africa are known. Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Sphyracephala beccarii was listed by Bezzi (1922) and beccarii (Fig. 1) has the most widespread distribu- Vaillant (1953) for El-Kantara in northern Algeria tion of the diopsids in the peninsula, extending into (not southern Algeria as mentioned by Vaillant). what is here considered as the Palaearctic area of the Ebrahim (2009) recorded for Egypt “D. apicalis”, a United Arab Emirates. The rather limited viewpoint misidentification forD. mayae. The flies were col- based on the distribution of Diopsidae tends to sup- lected in 1963 in Armant, near Luxor, not in the port the view that the Afrotropical Region in the more northern Assiut as stated by Ebrahim. Arabian Peninsula consists of the mountainous ar- A regularly returning discussion in papers on eas of southwest Saudi Arabia, Yemen, and southern the Fauna of the Arabian Peninsula concerns the Oman. The distribution of S. beccarii in the penin- delimitation of the Afrotropical and Palaearctic Re- sula confirms its status as the most widespread of all gions (e.g. Roselaar 2006, Kirk-Spriggs & McGregor Diopsidae, extending from Algeria to South Africa 2009, El-Hawagry et al. 2013). Sclater (1858) and and from Madagascar to the United Arab Emirates. Wallace (1876) identified the Palaearctic and Ethio- Hennig’s (1941) doubt concerning D. arabica “Es pian (later named Afrotropical) Regions. A transi- ist daher nicht sicher, ob er im paläarktischen oder tional Eremic or Saharo-Sindian Region was later äthiopischen Teil Arabiens gefangen wurde” can now also proposed. Wallace defined as border between be solved: this species only occurs in the Afrotropical the Palaearctic and Ethiopian regions in the Arabian part of the Arab Peninsula. Peninsula the Tropic of Cancer (~23°26’N). Later authors varied this border between 21°N and 28°N (see Roselaar 2006). For practical reasons, Crosskey Description of a cluster of S. beccarii flies (1980) used for his catalogue of Afrotropical Diptera in Wadi Darbat, Oman as border of the Afrotropical Region in the penin- On 10.ix.2014, 11 am, while birding in Wadi Darbat sula the northern boundaries of the modern state of (altitude 206 m) in the southern Sultanate of Oman, Yemen. For birds, Martins & Hirschfeld (1994) in- S. & M. Kühn, T. Wulf and R. Martin discovered a cluded the entire peninsula in the Palaearctic Region, cluster of S. beccarii flies. The cluster was on the bark with the exception of the mountains of southwest of live tree trunks next to the river and at a height Saudi Arabia, Yemen, and southern Oman. Roselaar of 0.5–3 m on the shaded side of the tree. The wadi (2006) proposed for the peninsula a revised southern itself is rocky with partly steep slopes and cliffs. The boundary of the Palaearctic Region which included site (Fig. 16) is sheltered from the wind and thanks the entire peninsula except the Afrotropical Moun- to the river more humid than the surrounding areas. tains of the south and southwest and the Oriental The air temperature was around 35°C. coastal plain of northern Oman. Based on studies of The fly cluster Figs( 17, 18, 20) consisted of one Diptera, Kirk-Spriggs & McGregor (2009) indicated major and about ten smaller clusters. Addition- that Wallace’s concept of the extent of the Afrotropi- ally, many flies were well spread over the trunk. At cal Arabian Peninsula is more accurate than Cross- the edge of the clusters few single S. beccarii could key’s concept of Yemen alone. Based on various Dip- be found. An area of about 35 dm2 was occupied tera families they supported the view that northern very densely with more than 15 flies per cm2. These Oman is the easternmost limit of the Afrotropical densely covered areas counted several layers of flies Region. El-Hawagry et al. (2013, 2015) studied a (Fig. 20). In addition, an area of at least 1.4 m2 was large set of insect orders in Al-Baha Province in Sau- covered with more than two flies per cm2. Based on di Arabia and concluded that this province ought to these numbers there were at least 52,500 flies in the be considered part of the Afrotropical Region rather densely occupied areas and about 28,000 flies in the than the Palaearctic Region. more sparsely covered areas. Altogether this leads to The distribution of the six Diopsidae species in a total of at least 80,500 specimens of S. beccarii. No the Arabian Peninsula is presented in Figs 1–4. Four particular smell was noted around the cluster. of these six species (S. beccarii, C. meigenii, D. ae The flies were walking around in the clusters, thiopica and D. mayae) belong to the small group of therefore the clusters were changing shape and, like- very common diopsids with an extensive distribu- wise the numbers of flies in the various aggregations tion in Sub-Saharan Africa, while at least one more were changing during our observation. Few flies were species (D. arabica) probably also occurs in Eritrea. seen flying around. When approached carefully, the S. beccarii is known from Algeria and D. mayae from flies didn’t react. However, when we made quick Egypt, both Palaearctic countries. The distribution movements, they flew up. Neither in the field nor on of the Diopsidae in the Arabian Peninsula concen- the photos, was any sign of intraspecific aggression trates on the mainly mountainous region from Al- noted. The cluster consisted of males and females, Baha Province in Saudi Arabia to the southern tip of but no mating behaviour was observed. Close to Yemen. In addition, C. meigenii (Fig. 2) also occurs the cluster a solitary C. meigenii was observed in the south-western tip of Oman. Sphyracephala (Figs 18, 19). Downloaded from Brill.com10/06/2021 02:46:57PM via free access
A review of clustering in Diopsidae at altitudes of 100–300 metres. One exception was Large to huge aggregations of Diopsidae are very formed by a cluster in early June but that was at common in the Afrotropical Region, especially in the around 1500 m and under very dry conditions. All dry season. They can be found along rivers or lakes cases concerned secluded positions: ceiling of cul- and in swampy areas or rain forests. These aggrega- verts, among roots overhanging drains, bushes over- tions can count thousands of specimens and can in- hanging a small stream and the shady face of a rock clude 10–20 species (Feijen, 1989 and later personal overhanging a pool. Sen observed: “On passing a net observations). In the Oriental Region large aggrega- over them they rose with a roar like a swarm of bees tions appear to be much less common, although they and a solid mass of several handsful (sic!) of flies was have sometimes been reported (Westwood 1837a, got in the net. Apparently the flies were originally Koningsberger 1915). congregated in a mass several individuals (? several A different issue in Diopsidae is the formation inches) deep. The mass consisted of flies of both sex- of clusters. The differences between clusters and es, in approximately equal numbers.” Mathur (1957) aggregations can be pointed out to be that clusters found “large congregations” of S. hearseiana in win- are much denser and can count tens of thousands of tertime (November–February). Mathur remarked normally conspecific flies. Occasionally a few speci- “Each cluster appears as a large compact black mass mens of another species can occur in a cluster, while of spot on the wall” and “They are readily disturbed only a few cases of really mixed clusters are known with one’s breath, and quickly fly about and settle on (see below). As the defining character of clusters the anything near-about”. The clusters were calculated formation of real layers of flies should be indicated (“estimated”) to be composed of between 4,762 and (Figs 20, 21). Flies then sit in a haphazard, random 50,952 specimens. way on top of each other. In the Nearctic Region, Flint (1956), Lavigne Kavanaugh (1977), working on carabid beetles, (1962) and Hochberg Stasny (1985) reported on later supported by Arnaud (1983), describing the clusters of S. brevicornis (Say) at their overwinter- phenomenon in a kelp fly, listed as possible mecha- ing site. These authors did not distinguish between nisms for the formation of aggregations: S. brevicornis and S. subbifasciata Fitch, so the latter species might also have been involved. Flint found clusters in a “small crack between two large blocks” 1. “Independent, individual response to an environ- and remarked “Undoubtedly there were thousands mental gradient (or gradients) leading to aggrega- present” and “over 150 were taken with a small kill- tion in an environmentally (abiotically) optimum ing vial before they all flew away”. Lavigne stated location.” that “Mating occurs almost immediately after the 2. “Individual response to some stimulus (or stim- adults come out of hibernation, while they are still uli) provided by other individuals, leading to ag- clustered at the overwintering site”. gregation at a common location.” In Algeria, in July 1949 Vaillant (1953) found 3. “A combination of the first two.” among rocks in an oasis “de véritables essaims de Sphyracephala Beccarii”. Feijen (1984) found in the The first mechanism could also be described as a dry season (22.viii.1974, Diampwe River, 1152 m) more passive system, leading to “non-intended” ag- in Malawi among rocks in a river bedding a dense gregations, while in the second one an active system mass of S. beccarii. After being disturbed they flew leads to aggregations. In Diopsidae the first mecha- up in a dense cloud. A single sweep of the net yielded nism could account for aggregations as found in the more than 6,000 specimens, the size of the whole dry season in Africa, while the second mechanism mass estimated to be in the order of 100,000 speci- appears more likely in the formation of clusters. mens. The percentage ♀♀ came to 47%. Reports on clusters in the Diopsidae all refer to Reports on cluster formation in other Diopsidae species of Sphyracephala. Brunetti (1907, 1919) genera are rare. Marion Kotrba (pers. comm.) ob- was the first one to mention “vast numbers” for served cluster formation in Howick (South Africa) in Sphyracephala hearseiana (Westwood). He recorded early May 1992. This involved Diopsis mayae and a these Indian flies “in profusion under a low arch species of the Diopsis atricapilla species group. These over a roadside ditch in Cawnpore” on or around flies already formed dense aggregations during the 30.xi.1904. Furthermore, he mentioned these flies day. At late dusk, they flew away under a bush. There, were “clustered very thickly together on the inside Kotrba found them in the early morning gathered walls of the ground floor of that deserted building” on a single, kinked stem of grass, on the part that of the old Residency at Lucknow on 4.xii.1904. Sen was hanging down. The cluster was really dense and (1922) described a number of additional clusters for several flies deep, like a bee swarm. When she tried S. hearseiana and also provided the first photograph to collect them, she touched the grass and all flies of a cluster. All observations were in wintertime and were up and flying within an instant. Martin Grimm Downloaded from Brill.com10/06/2021 02:46:57PM via free access
(pers. comm.) observed and photographed clus- In the Oriental Region a common, but not re- ters in Matema, Tanzania, on 6.vi.2014 at around lated, aggregation phenomenon occurs in the diop- 11 o’clock a.m. The clusters (Fig. 21) mainly consist- sid genera Teleopsis Rondani and Cyrtodiopsis Frey. ed of Dia. varians, but in between were also S. mun These aggregations have been referred to as sleeping roi Curran and a few S. beccarii. The clusters were on communities or societies, nocturnal clusters, noctur- leaves around 1.5 m above a small montane river in nal lekking aggregations, harems and aggregations the forest. The place was somewhat cooler than the on nocturnal roosting sites (de la Motte & Burkhardt further ambient temperature of 28–30°. In Maputo, 1983, Wilkinson & Dodson 1997, Cotton et al. Mozambique, the first and third author several times 2015). Typically, flies gather at dusk in relatively found clusters of C. meigenii on lower leaves of gar- small conspecific aggregations on thread-like hang- den shrubs. This was in the cold season (June–July) ing vegetation. Males engage in ritualized fights to and involved several thousands of flies. A curious ad win a harem (de la Motte & Burkhardt 1983), while hoc cluster was once observed in Malawi by the first “The vast majority of matings occur in these ag- and third author. Very early in the morning, on an gregations during the dawn and dusk period, when unusually cold day, they visited a rice scheme to col- males attempt to mate with females in their harem” lect samples of rice hills. While changing into rubber (Cotton et al. 2015). boots, they left their shoes on a paddy bund. Re- turning some 30 minutes later, they noted that their shoes were covered by layers of Diopsis longicornis Acknowledgements Macquart, the well-known rice stem-borer. The only Martin & Susanne Kühn made two of the Oman explanation could be that the flies were attracted to photographs available, while Martin Grimm sup- the warm temperature of the shoes. This phenom- plied the photograph of a Diasemopsis cluster in enon was never observed again during many years of Tanzania. We are grateful to Martin Hauser, Martin study of D. longicornis. The only other occasion in- Grimm and Marion Kotrba for discussions about volving this species was on 26.vi.1974 at the Khan- cluster formation. Daniel Whitmore (BMNH) and da River (Chilwa Plain, Malawi, 675 m) when the Eliane De Coninck (MRAC) provided access to the same observers noted a dense cluster of thou- Diopsidae collection of their museums. The original sands of D. longicornis and D. mayae in a clump of idea for a paper on Arabian Diopsidae came from reeds. At the same location, aggregations were regu- Prof. Dr William Büttiker-Otto (1921–2009), larly observed, but only this once a real cluster was who proposed it to the first author in 1989. The involved. comments by anonymous referees are greatly It appears certain that real clusters of diopsids, appreciated. involving massive numbers of up to 100,000 speci- mens, can only be the result of the second mechanism stated by Kavanaugh (1977): individual response to References some stimulus (or stimuli) provided by other indi- Anagnou-Veroniki, M., P. Papaioannou-Souliotis, E. viduals. Initial aggregation in an environmentally Karanastasi & C.N. Giannopolitis, 2008. New records optimum location can play a part just as well. The of plant pests and weeds in Greece, 1990–2007. – phenomenon of cluster formation, especially in Hellenic Plant Protection Journal 1: 55–78. Sphyracephala, appears clearly linked to the process Arnaud, P.H., 1983. Aggregation of Coelopa (Neocoelopa) of hibernation or estivation. In clusters, mating be- vanduzeei Cresson on the Monterey Peninsula Coast, haviour has never been observed. Lavigne (1962) ob- California, and notes on the family (Diptera: Coelopi- served that mating occurs almost immediately after dae). – The Pan-Pacific Entomologist 58: 245–249. the hibernation period. It appears to be a common Bezzi, M., 1908a. Diagnoses d’espèces nouvelles de Dip- character for Diopsidae clusters that, when even su- tera d’Afrique. – Annales de la Société Entomologique de Belgique 52: 374–388. perficially disturbed, the flies can immediately fly up Bezzi, M., 1908b. Ditteri Eritrei raccolti dal Dott. Andreini en masse. This is the more striking as it happens also e dal Prof. Tellini. II. Diptera Cyclorrhapha. – Bollet- during the estivation or hibernation period with as- tino della Società Entomologica Italiana 39: 3–199. sumed lower rates of metabolism. It seems likely that Bezzi, M., 1922. Note sur la présence en Algérie du Sphyra the explanation for the occurrence of multi-layer cephala Hearseiana Westw. de l’Inde et sur la synony- clusters in Diopsidae must be sought in the direc- mie de ce Diptère. – Bulletin du Muséum National tion of maintenance of an optimal micro-habitat. d’Histoire Naturelle 28: 69–72. The maintenance of a certain level of temperature Bruggen, A.C. van, 1961. Diptera (Brachycera): Diopsi- and perhaps also humidity, would then enable the dae. A partial revision of the Diopsidae or stalk-eyed flies to react very fast to disturbance. The importance flies of Southern Africa. – In: B. Hanström, P. Brinck of temperature is clearly shown by the behaviour of & G. Rudebeck (Eds), South African Animal Life, Re- D. longicornis on the bund in Malawi. sults of the Lund University Expedition in 1950–1951, Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Vol. 8, pp. 415–439. Almqvist & Wiksells Boktryckeri Eggers, F.O., 1916. On some new and incompletely AB, Uppsala, Sweden. known species of the family Diopsidae from British Brunetti, E., 1907. XII.–Notes on Oriental Diptera. I.– East Africa. – In: V.A. Dogiel (Ed.), Scientific results Note on Sphyracephala hearseyana Westwood, with a of the zoological expedition to British East Africa and list of the Oriental species of Diopsinae. – Records of Uganda, made by Prof. V. Dogiel and I. Sokolow in the Indian Museum 1: 163–166. the year 1914, vol. I, no. 6, pp. 3–32, unnumbered pl., Brunetti, E., 1919. Review of progress in our knowledge of Russian on pp. 3–20, English on pp. 21–32. Petrograd, Oriental Diptera during the last two decades. – Jour- Russia. nal of the Asiatic Society of Bengal (New Series) 14: Eggers, F.O., 1925. Diopsiden aus Deutsch-Ostafrika, mit 358–371. einem Nachwort über die Stielaugen der Diopsiden. – Brunetti, E., 1926a. Notes on some Belgian Congo Dip- Zoologische Jahrbücher 49: 469–500, pl. 6. tera. – Revue Zoologique Africaine 13: 165–173. El-Hawagry, M.S., M.W. Khalil, M.R. Sharaf, H.H. Fadl Brunetti, E., 1926b. New species of Diopsidae (Diptera) & A.S. Aldawood, 2013. A preliminary study on the from the Belgian Congo. – Revue Zoologique Africaine insect fauna of Al-Baha Province, Saudi Arabia, with 14: 73–84. descriptions of two new species. – ZooKeys 274: 1–88. Brunetti, E., 1928. Revisionary notes on the Diopsidae El-Hawagry, M.S., M.R. Sharaf, H.M. Al Dhafer, H.H. (Diptera). – Annals and Magazine of Natural History, Fadl & A.S. Aldawood, 2015. Addenda to the insect Ser. 10, 2: 266–275. fauna of Al-Baha Province Saudi Arabia. – Journal of Carr, M., S. Cotton, M. Földvári & Kotrba, M., 2006. A Natural History 50: 1209–1236. description of a new species of Diasemopsis (Diptera, El-Hawagry, M.S., M.S. Abdel-Dayem, A.A. Elgharbawy Diopsidae) from the Comoro Islands with morpholog- & H.M. Dhafer, 2016. A preliminary account of the ical, molecular and allometric data. – Zootaxa 1211: fly fauna in Jabal Shada al-A’la Nature Reserve, Saudi 1–19. Arabia, with new records and biogeographical remarks Cogan, B.H. & J.F. Shillito, 1980. 47. Family Diopsidae. (Diptera, Insecta). – ZooKeys 636: 107–139. – In: R.W. Crosskey (Ed.), Catalogue of the Diptera of Feijen, H.R., 1978. Diopsidae (Diptera: Acalyptratae) from the Afrotropical region, pp. 583–587. British Museum Togo and Zaïre. – Stuttgarter Beiträge zur Naturkunde (Natural History), London, UK. 318: 1–25. Collart, A., 1954. Les Sphyracephala du Congo Belge (Dip- Feijen, H.R., 1984. Studies on the systematics, ecology and tera Diopsidae). Miscellanea Zoologica H. Schouteden. economic importance of the Diopsioinea (Diptera). – – Annales du Musée Royal du Congo Belge, Tervuren PhD dissertation, University of Leiden. H.M. Ooster- (Belgique), nouvelle série in-4o. Sciences Zoologiques huis, Leiden, The Netherlands. 1: 329–331. Feijen, H.R., 1987. A revision of the Diopsidae (Diptera) Cotton, A.J., S. Cotton, J. Small & A. Pomiankowski, described by J.W. Dalman. – Entomologica Scandi- 2015. Male mate preference for female eyespan and navica 17: 409–422. fecundity in the stalk-eyed fly, Teleopsis dalmanni. – Feijen, H.R., 1989. Diopsidae. Cyclorrhapha III Behavioral Ecology 26: 376–385. (Schizophora other than Calyptratae). – In: G.C.D. Crosskey, R.W. (Ed.), 1980. Catalogue of the Diptera of Griffiths (Ed.), Flies of the Nearctic Region, Vol. 9, part the Afrotropical Region. – British Museum (Natural 12, pp. 1–122. E. Schweizerbart, Stuttgart, Germany. History), London, UK, [iv] + 1437 pp. Feijen, H.R. & C. Feijen, 2009. Diopsis (Diopsidae, Dip- Curran, C.H., 1928. Review of the African species of tera) with unusual wing spots: Two new species from Sphyracephala Say. – Annals and Magazine of Natural Malawi with a longer eye span in females than in males. History 10: 274–275. – Zoologische Mededelingen Leiden 83: 701–722. Curran, C.H., 1931. Descriptions of new species of Dia Feijen, H.R. & C. Feijen, 2012. A new species of Diopsis L. semopsis Rondani (Diopsidae, Diptera). – American (Diptera: Diopsidae) from South Africa and Swaziland, Museum Novitates 463: 1–17. and brief review of African species with a large apical Dalman, J.W., 1817. Anmärkningar vid slägtet Diop- wing spot. – African Invertebrates 53: 125–142. sis, jemte beskrifningar och teckningar på trenne Feijen, H.R. & C. Feijen, 2017. Diopsis mayae sp. n. (Dip- dithörande nya arter. – Kongliga Vetenskaps Acad- tera, Diopsidae), a dominant stalk-eyed fly, occurring emiens Handlingar 38: 211–219. from South Africa to Saudi Arabia. – Tijdschrift voor Dawah, H.A. & M.A. Abdullah, 2008. First records of Entomologie 160: 61–74. the stalk-eyed flies (Diopsidae: Cyclorrhapha), a pest Flint, O.S., 1956. Hibernation of the diopsid fly,Sphyra of Poaceae, from South-Western Saudi Arabia. – King cephala brevicornis Say. – Bulletin of the Brooklyn En- Khalid University Journal of Natural, Medical, and tomological Society 51: 44. Technical Sciences 4: 86–95. Frey, R., 1921. Studien über den Bau des Mundes der nie- de la Motte, I. & D. Burkhardt, 1983. Portrait of an Asian deren Diptera Schizophora nebst Bemerkungen über stalk-eyed fly. – Naturwissenschaften 70: 451–461. die Systematik dieser Dipterengruppe. – Acta Societatis Ebrahim, A.M., 2009. Described a new recorded family pro Fauna et Flora Fennica 48: 1–245. Diopsidae of (Order Diptera) with its species Diopsis Guiglia, D., 1957. I tipi di Ditteri di Camillo Rondani e di apicalis in Egypt. – Egyptian Academic Journal of Bio- altri autori appartenenti alle collezioni del Museo Civ- logical Sciences A. Entomology 2: 135–145. ico di Storia Naturale di Genova. – Annali del Museo Downloaded from Brill.com10/06/2021 02:46:57PM via free access
Civico di Storia Naturale Giacomo Doria, Genova 69: Rohlfien, K. & B. Ewald, 1970. Katalog der in den Samm 193–207. lungen des ehemaligen Deutschen Entomologischen Hauser, M., J.C. Deeming & J.H. Stuke, 2011. Order Institutes aufbewahrten Typen. VIII (Diptera: Cyclor- Diptera, family Diopsidae. – In: A. van Harten (Ed.), rhapha: Schizophora: Acalyptratae). – Beiträge zur En- Arthropod fauna of the United Arab Emirates, Vol. 4, tomologie 22: 407–469. pp. 765–768. Dar Al Ummah, Abu Dhabi. Rondani, C., 1873. Muscaria exotica Musei Civici Janu- Hendel, F., 1923. Afrikanische Diopsiden (Diptera). – ensis observata et distincta. Fragmentum I. Species Wiener Entomologische Zeitung 40: 33–42. aliquae in Abyssinia (Regione Bogos), lectae a Doct. Hennig, W., 1941. 39c Diopsidae. – In: E. Lindner (Ed.), O. Beccari et March. O. Antinori, anno 1870–71. – Die Fliegen der palaearktischen Region: 5(7), Lieferung Annali del Museo Civico di Storia Naturale Giacoma 143, pp. 1–8. E. Schweizerbart, Stuttgart, Germany. Doria, Genova 4: 282–294. Hochberg Stasny, T.A., 1985. Biology, behavior and life Rondani, C., 1875. Muscaria exotica Musei Civici Janu- cycle of Sphyracephala brevicornis Say (Diptera: Diopsi- ensis observata et distincta. Fragmentum III. Species dae). – MSc thesis, West Virginia University, Morgan- in Insula Bonae fortunae (Borneo), Provincia Sarawak, town, VA. USA. annis 1865–68, lectae a March. J. Doria et Doct. O. Karsch, F., 1888. Bericht über die durch Herrn Lieutenant Beccari. – Annali del Museo Civico di Storia Naturale Dr. Carl Wilhelm Schmidt in Ost-Afrika gesammelten Giacoma Doria, Genova 7: 421–464. und von der zoologischen Abtheilung des Königli- Roselaar, C.S., 2006. The boundaries of the Palearctic re- chen Museums für Naturkunde in Berlin erworbenen gion. – British Birds 99: 602–618. Diptera. – Berliner Entomologische Zeitschrift 31: Sclater, P.L., 1858. On the general geographical distri- 367–382, pl. 4. bution of the members of the class Aves. – Journal Kavanaugh, D.H., 1977. An example of aggregation in and Proceedings of the Linnean Society (Zoology) 2: the Scaphinotus subgenus Brennus Motschulsky (Cole- 130–145. optera: Carabidae: Cychrini). – The Pan-Pacific Ento- Séguy, E., 1949. Diopsides de Madagascar. – Memoires de mologist 53: 27–31. l’Institut Scientifique de Madagascar A 3: 65–76. Kirk-Spriggs, A.H. & G. McGregor, 2009. Disjunctions in Séguy, E., 1955. Diptères Diopsides africains nouveaux ou the Diptera (Insecta) fauna of the Mediterranean Prov- peu connus. – Bulletin de l’Institut Français d’Afrique ince and southern Africa and a discussion of biogeo- Noire Sér. A, Sciences Naturelles 17: 1102–1124. graphical considerations. – Transactions of the Royal Sen, S.K., 1922. Life-histories of Indian insects. Diptera: Society of South Africa 64: 32–52. Sphryracephala (sic!) hearseiana, Westw. – Memoirs of Koningsberger, J.C., 1915. Java, zoölogisch en biologisch. the Department of Agriculture in India (Entomological – Departement van Landbouw, Nijverheid en Handel, Series) 7(6): 33–38. Buitenzorg, Nederlands Indië, 663 pp. Shillito, J.F., 1971. The genera of Diopsidae (Insecta: Dip- Kotrba, M., 2009. Prosphyracephala kerneggeri spec. nov. tera). – Zoological Journal of the Linnean Society 50: (Diptera: Diopsidae). A new stalk-eyed fly from Baltic 287–295. amber (Diptera, Diopsidae). – Spixiana 32: 187–192. Steyskal, G., 1972. A catalogue of species and key to the Lavigne, R., 1962. Immature stages of the stalk-eyed fly, genera of the family Diopsidae (Diptera: Acalyptratae). Sphyracephala brevicornis (Say) (Diptera: Diopsidae) – Stuttgarter Beiträge zur Naturkunde, Serie A (Biolo- with observations on its biology. – Bulletin of the gie) 234: 1–20. Brooklyn Entomological Society 57: 5–14. Vaillant, F., 1953. A propos d’un Diopside d’Algérie Lindner, E., 1962. Studien an afrikanischen Diopsiden Sphyracephala Beccarii Rondani. – L’Entomologiste 9: (Dipt.). – Stuttgarter Beiträge zur Naturkunde 94: 11–12. 1–18. Wallace, A.R., 1876. The geographical distribution of ani- Macquart, J., 1843. Diptères exotiques nouveaux ou peu mals, Vol. 1. – MacMillan & Co, London, UK, xxi + connus. Tome deuxième. 3e partie. – Memoires de la 503 pp. Société Royale des Sciences, de l’Agriculture et des Arts Westwood, J.O., 1837a. On Diopsis, a genus of dipter- de Lille [1842]: 162–460, pls 1–36. ous insects, with descriptions of twenty-one species. Martins, R.P. & E. Hirschfeld, 1994.Where are the limits – Transactions of the Linnean Society of London 17: of the Western Palearctic? – Bulletin of the British Or- 283–313, pl. 9. nithologists’ Club 114: 207–208. Westwood, J.O., 1837b. Descriptions of some new spe- Mathur, R.N., 1957. Large clusters of SphyracephaIa hear cies of Diopsis. – Transactions of the Linnean Society of seiana Westwood (Diptera: Diopsidae) at New Dehli. London 17: 543–550, pl. 28. – Current Science (Bangalore) 26: 183–184. Wilkinson, G.S. & G.N. Dodson, 1997. Function and Ôhara, K., 1993. First record of the family Diopsidae from evolution of antlers and eye stalks in flies. – In: J. Choe Japan (Diptera, Brachycera). – Bulletin of the Tokushi- & B. Crespi (Eds), The evolution of mating systems in ma Prefectural Museum 3: 39–47. insects and arachnids, pp. 310–328. Cambridge Uni- Osten Sacken, C.R., 1882. Diptera from the Philippine versity Press, Cambridge, UK. Islands, brought home by Carl Semper. – Berliner En- tomologische Zeitschrift 26: 83–120, 187–252.
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