Diptera: Fam. Chirononlidae of the Arabian Peninsula - Entomology

FAUNA OF SAUDI ARABIA la,
INTRODUCTION
The Chironomidae are an abundant and widespread family of flies, often known as non-biting
midges. As the popular name implies, unlike their relatives, the Chironomidae do not bite. However,
they are implicated in human disease when nuisance numbers of swarming adults are unavoidably
inhaled, thereby inducing allergic disease in susceptible individuals (CRANSTON et al. 1981, 1983).
In their behaviour, adult chironomids are evident when swarming, particularly at dusk near water,
the site of the development of the immature stages. Larvae, often possessing haemoglobin pigments
and known as "bloodworms", may reach densities of 1 00 000 m- 2 in polluted habitats such as sewage
settlement lagoons (e. g. Jeddah sewage works). However, chironomid larvae are not restricted to polluted
habitats but a wide range of species occur in all aquatic environments, including Arabian wadis
and oases. This broad ecological tolerance and diversity of species means that Chironomidae have
been and remain the subject of much study in relation to their utility as biological indicators of water
quality, as reviewed by PL"DER (1986).
In much of Europe and eastern North America, the taxonomy and ecological requirements of
Chironomidae are well known. Elsewhere, including the southern Palaearctic, much basic study is still
required. REISS (1977) observed the lacunae in our knowledge of the Palaearctic fauna and drew particular
attention to the poorly known eastern Mediterranean, an impediment to discussion of regional
chironomid biogeogeography. REISS himself contributed to the resolution of this problem in his treatment
of the chironomid faunas of Turkey (REISS 1985) and Syria (REISS 1986). The first documentation
of any Chironomidae from Saudi Arabia appears to be CRANSTON'S (1989) description of nine
new species from south-western Asia, including Saudi Arabia.
In this paper we record all species of Chironomidae we have seen from the Arabian peninsula.
Species discovered are not redescribed fully in the current mode, partially following CRANSTON &
OLIVER'S (1988) diatribe against lengthy descriptions in chironomid systematics, but keys to subfamilies,
genera and species are given accurate means of identification in other works referenced, discriminatory
characters discussed and illustrations provided of the hypopygia to allow identification of
adult male Arabian Chironomidae. No assessment of the phylogenetic relationships is made although
we believe that in world-wide revisions cladistic studies are an essential precursor to historical biogeographical
analysis. Our discussion on biogeography is limited by our lack of detailed knowledge
of sister group relationships for taxa studied, and therefore relies upon examination of the present
day ranges of the fauna, followed by a search for congruent patterns in other organisms, and a correlated
earth history.
MATERIALS AND METHODS
Chironomidae were collected by sticky trap in Saudi Arabia by W. BUttiker, or by light trap in Oman
by R.P. Lane. As discussed in CRANSTON (1989), sticky trap material temporarily preserved in Berlese
mountant retains castor oil and is inadequate for description, and often for recognition. Berlese
mountant (and Hoyer's and related Chloral Hydrate based mountants) is unsuitable for permanent
preservation of slide mounted Diptera through variable, but probable eventually universal crystallisation
or other opacity (contra DISNEY & HENSHAW, 1988). CRi\NSTON (1989) discusses the recovery and
appropriate method for making permanent mounts in Euparal.
Descriptions and morphological terminology follow SAETHER (1980). Generic placements follow
the keys and diagnoses for adults of Holarctic Chironomidae to be published in WlEDERHOLM (1989).
In illustrations of the hypopygia, the left side is shown in dorsal view; the more schematic,
slightly detached, right half is a portrayal of the ventral view, with underlying structures such as the
237

238 P. S. CRA:'JSTON, D. D. JUDD
apodemes shown hatched. Setation and complete distribution of microtrichia is of restricted illustration
on the right (ventral) half.
Geographical localities within Saudi Arabia follow a north-west to south-east sequence of provinces,
listed as follows: Jawf, Tabuk, Hail, Medina, Gasim, Makkah, Riyadh, Eastern Province, Baha,
Asir, Najran, Gizan. Distributional information for Afrotropical taxa comes from FREEMAN & CRAN­
STON (1980), supplemented by DEjoCX (1984) for Guinea, HARRlSOK (1987) for Ethiopia and HARE &
CARTER (1987) for Nigeria. Palaearctic distributional information is derived from CRANSTON & ASHE
(in press) and Oriental distributions partially follow CHAUDHURI & GUHA (1987).
Unless stated otherwise, all specimens are mounted in Euparal and deposited in the Diptera section
of the Entomology Department, British Museum (Natural History), London (BMNH). Duplicate
representatives of as many Arabian species as possible are deposited in NHMB and ZSM.
Abbreviations of repositories in the text are as follows:
ANIC: Australian National Insect Collection, Division of Entomology, CSIRO, Canberra, Australia.
BMNH: Entomology Department, British Museum (Natural History), London, U.K.
MRAC: Entomologie, Musee Royal de l'Afrique Centrale, Tervuren, Belgium.
NHMB: Entomologie, Naturhistorisches Museum, Basle, Switzerland.
SMNH: Naturhistoriska Riksmuseet, Stockholm, Sweden.
ZSM: Zoologische Staatssammlung, Munich, German Federal Republic.
Other abbreviations used in text:
i.v.: inferior volsella (of male hypopygium)
m o.d.: altitude in metres over datum.
m.v.: median volsella (of male hypopygium)
S.V.: superior volsella (of male hypopygium)
Key to subfamilies of Saudi Arabian Chironomidae
1. Wing with crossvein MCu present (figs 1,2) TanypoJinae
Wing lacking crossvein M Cu ............................... . 2
2. Foreleg with tibia longer than tarsomere 1 (Leg Ratio < 1). Hypopygium with gonosty­
Ius articulating with gonocoxite, such that gonostylus is variably incurved with apical
megaseta directed anteriorly OrthociaJiinae
Foreleg with tibia shorter than tarsomere 1 (Leg Ratio > 1). Hypopygium with gonostylus
rigidly affixed to gonocoxite, if articulating with gonocoxite (Stictochironomus)
gonostylus lacks megaseta and may be strongly incurved Chironominae
Subfam. Tanypodinae
Key to species of Saudi Arabian Tanypodinae
1. Wing cross vein MCu clearly proximal to FCu (fig. 1) .................. . 2
Wing cross vein MCu ends at FCu (fig. 2) .. ...................... . 4
2. Wing membrane bare. Distinctive pale body colour with yellow to red vittae and black
median scutal stripe, white legs with black distal tarsomeres, white abdomen with median
longitudinal black stripe. Outer heel of gonocoxite not developed (fig. 5)
ProciaJius (PsiJotanypus) reiJi

240 P. S CRANSTON. D. D. JUDD
tinctive by virtue of the restriction of the wing macrotrichia to the apical 114 of cell r4+5 and sometimes
the extreme apex of m1+ 2 ' Representatives of all other western Palaearctic and Afrotropical
species have macrotrichia much more widely distributed over the wing membrane than in P. apicalis
sensu FREEMAN (I955b). Although the preservation of Saudi Arabian specimens has led to loss of most
wing macrotrichia, it is possible to detect the restricted distribution through observation of remnant
pits. Furthermore, although fading of any wing pattern in many specimens has occurred, it is unlikely
that the strong patterns of related Procladius species would have been completely lost.
Although the taxonomic decision to include all species with reduced wing macrotrichia is substantiated
by wing, body and leg colour, there is rather extensive hypopygial variation. The relative
length of the phallapodeme varies between about 114 up to half gonocoxite length in some specimens.
The shape of strut 2 (ROBACK 1971) (homologised with the superior volsella by SAETHER [1980]) varies
with orientation, but there seem to be genuine differences in length and apicomedial construction.
Finally, the outer "heel" of the gonostylus is variably developed, but such variation, although associated
partially with orientation, seems to be widespread within the genus.
Additional doubt concerning identity and species limits comes indirectly from the distribution: P.
apicalis is virtually alone amongst the Afrotropical species present in Arabia that occur as far north as
Jawf and Hail. In these regions the fauna is virtually exclusively Palaearctic (see Biogeographic section
of Discussion).
Whether there is more than one species, and if they are truly P. apicalis will only be confirmed
with the assistance of associated immature stages, particularly the pupa.
Although FREEMAN (1955b) cites the antennal ratio (A.R.) as being nearer I (compared to P. brevipetio!atus),
recalculation from newly slide mounted specimens examined by FREEI,tAN (1955b) show
an A.R. of 1.8-2.0, i. e. not allowing discrimination from brevipetiolatus.
Proc1adius (Holotanypus) brevipetiolatus (Goetghebuer, 1935)
Trichotanypus brevipetioiatus Goetghebuer, 1935. Rev. Zool. Bot. Afr. 27: 355. Type-locality: Belgian Congo (Zaire),
Pare National Alben, Vitshumbi [Type(s) not seen].
Prodadius brevipetioiatllS. Freeman 1955b; Bull. Br. Mus. Nat. Hist. Ent. 4: 56.
Material: 1

242 P. S. CRANSTO"l, D. D. JUDD
Dis tri bu tion: Afrotropical region: Guinea (DEJOUX 1984), Sudan. This constitutes the first record
from outside the Afrotropical region.
Comments: Procladius (Psiiotanypus) reidi was described from Adok and Melut on the White
.Nile in central Sudan, from females alone. Although DEJoLx (1984) reported a male from a tributary
of the River Niger in Guinea, the male has not been described previously. Females from Saudi Arabia
are identical with those from Sudan; males have a similar, distinctive colouration and are clearly conspecific
with females from Sudan and Saudi Arabia.
Ablabesmyia (Ablabesmyia) longistyla Fittkau, 1962
Ablabesmyia longistyla Fittkau, 1962 Abh. Larvalsyst. Ins. 6: 436. Type-localities: various in Austria & Germany (Types
not examined).
Materia I: 10',299, Saudi Arabia: Makkah, Wadi Maraum, 22° 16'01 39° 44'E, 280 m o.d., 3.-4. V. 1984, W. Bilttiker. 1
0', Saudi Arabia: Makkah, Wadi Turabah, 20 c 29'N 41°15'E, 1400 m o.d., 21. IV. 1980, W.BUttiker. - 6 cJcJ (3 to NHMB),
Saudi Arabia: Riyadh, Ain al Burj, 22° 10'01 46" 42'E, 550 m o.d., 15. IX. 1981, W. Biittiker. - 1 0', Saudi Arabia: Eastern province,
Hofuf, 150 m o.d., 25°24'01 49°28'E, 28.V.1978, W.Biittiker. - 10', Saudi Arabia: Eastern province, Qatif, 26°33'N
49° 59'E, 8 m o.d., 18.-19. IX. 1979, W. BUttiker. 1 9, Saudi Arabia: Bab, Adnan, 20° 26':-.r 41' 31'E., 1350 m o.d., 21.-23. IX.
1978, W. Btittiker. - 1 C(, Saudi Arabia: Baha, Wadi Diyhan, 19° 48'N 41 c 36'[,830 m o.d., 7. III. 1984, W. Btittiker. - 1 cJ, Saudi
Arabia: Baha, Wadi lIyab, 20° 07'N 40° 57'E. 160 m o.d., 3. II. 1984, W. Btittiker. - 2 99, Saudi Arabia: Najran, Talaa, 18" C4'N
43° 57T, 1420 m o.d., 23.-24. IX. 1980, W. Btittiker. lOman: Tahwa near Sur, 'light-trap in goat pen', 25.XI. 1986, R. P.
Lane (B.MNH).
Leg (anterior femur, tarsus and tarsomere 1): fig. 3.
Hypopygium: figs. 6, 7.
Distribution: Widespread in the western Palaearctic, eastwards to Turkey, Lebanon and Syria.
A specimen from Egypt (Luxor, 12.1. 1981, P.S.Cranston, BM 1981-79) appears to be the first record
for this country.
Comments: Saudi Arabian A. longistyla lack a distinct dark mark at the apex of wing vein R2+3'
thus apparently differing from western European specimens. Since all specimens here examined are
variably faded and lack most dark macrotrichia towards the anterior margin and submargin of the
wing, it is not possible to determine if this dark area is truly lacking, or the result of fading and abrasion.
The identity of the male genitalia suggests con specificity with British specimens examined.
Ablabesmyia (Ablabesmyia) monilis (Linnaeus, 1758)
Tipula monilis Linnaeus, 1758. - Syst. :-.rat. 10: 587. Type-locality: 'Europe' (not examined).
Ablabesmyia monilis. Fittkau, 1962; Abh. Larvalsyst. Ins. 6: 437.
Materi al: I cJ, Saudi Arabia: ]awf, Al ]awf, 29° 52'N 39° 53,E, 610 m o.d., I.-LXI. 1986, W. Btittiker.
Hypopygium: fig. 8.
Distribution: Widespread, from Kearctic region (subarctic Canada and Alaska south to Carolinas),
Europe, Soviet Union, Japan, Korea and Taiwan.
Comments: Ablabesmyia monilis has a characteristic hypopygium. In this Saudi Arabian specimen
the anterior margin of the wing has distinct dark areas at the apices of Rp R2+3 and R4+s: the
absence of a distinct mark at the apex of R2+3 is used as the means of recognizing regional females of
Ablabesmyia as A. longistyla rather than A. monilis.
Conchapelopia trifascia (Freeman, 1954)
Pentaneura trifoscia Freeman, 1954. - Proc. Roy. Ent. Soc. Lond. (B) 23: 176. Type-locality: (South Africa) Cape Province,
Tulbagh Barrage (Holotype 9 examined, BMNH). Pentaneura trifascia. - Freeman, 1955b; Bull. Br. Mus. Nat. Hist. Ent.
4: 25.
Conchapelopia trifoscia. Harrison 1971; Can. Ent. 103: 387.
Material: 1 Saudi Arabia: Baha, Adnan, 1350 m o.d., 20026'N 41°3I'E, 21.-23.IX.1978, W.Buttiker (BM:-.rH).

244
Larsia rutsburuiemis (Goetghebuer, 1935)
P S, CRANSTON, D, D, JUDD
Ablabesmyia mtshHrHiensis Goetghebuer, 1935. Rev. Zool. Bot. Afr. 27: 362, Type-locality: Belgian Congo (= Zaire),
Rutshuru (Para type d examined, B,'\1NH).
Penlanellra rut5hllruiensis, - Freeman 1955; Bull. Br. Mus, Nat. Hist. Ent. 4: 27.
Larsia rutshumiensis. - Harrison 1971; Can, Ent. 103: 389.
Material: 2 dd, Saudi Arabia: Medina, Fare, 23°05'N 39°54'E, 1160 m o.d., 13.-14.XI.1986, W.Btittiker (I to
NHMB). 2 Saudi Arabia: Makkah, Wadi Maraum, 22" 16'N 39° 44'[,280 m o.d., 3,-4. V, 1984, W. Btittiker. I d, Saudi
Arabia: Makkah, Wadi Qust, 20 c 56'N 41 °06'£, 1400 m o.d., 28. II. 1984, W Btittiker. - 2 dd, Saudi Arabia: Baha, AI Mindak,
20° 07'N 41 ° 18'E, 1760 m o,d., 8, IV, 1980, W. Buttiker. 1 Saudi Arabia: Asir, AI Dalhan, 18°01'N 43° 24'E, 2180 m o.d.,
19.-20. IX. 1980, W. Btittiker.
Wing: fig. 2.
Thorax: fig. 10.
Hypopygium: fig. 1t.
Distribution: Afrotropical region; from South Africa, Zaire, Zimbabwe. This constitutes the
first record outside the Afrotropical region.
Comments: FITTKAU'S (1962) definition of Larsia was expanded by HARRISO,," (1971) to encompass
Afrotropical species, including L rutshuruiensis, with patterned wings. The presence of Larsiatype
tibial combs (although very small) in conjunction with the characteristic darkening of the cross
veins of the wing render this species distinctive.
Larsia teesdalei (Freeman, 1955)
PentaneHra teesdalei Freeman, 1955b. - Bull. Bf. Mus. Nat. Hist. Ent. 4: 26. Type-locality: Kenya, Kitui (Holotype d
examined, BJ'vlNH) ,
Larsia teesdalei. Harrison 1971; Can. Ent. 103: 389.
Material: I d, Saudi Arabia: Baha, Wadi Ilyab, 20 0 07'I\: 40° 57'E, 160 m o,d., 3.11.1984, W.Btittiker.
Distribution: Afrotropical region: Kenya and Zimbabwe. A specimen from Nigeria (N Nigeria,
Kano, fast stream below Tiga Dam, 20. II. 1981, A. Brown, BMNH) is the first record of the species
from this country. The present record is the first for the species from outside the Afrotropical region.
Comments: This solitary specimen belongs to the genus Larsia by virtue of the characteristic
Iyrate tibial spurs and scutal tubercle. The specimen is badly faded, yet the distinctive darkening of
the femoral apex and tibial base ("knee") together with the faint, but characteristic wing pattern, preclude
identity with any other described Larsia. The genitalia on the available specimen is too damaged
for illustration.
Paramerina vaillanti Fittkau, 1962
l'aramerina vaillanti Filtkau, 1962, - Abh. Larvalsyst. Ins. 6: 335. Type-locality: Algeria, Oasis Beni Abbes (Holotype d
examined, ZSM).
Material: 6 dd, Saudi Al'abia, Jawf, Al Jawf, 29 c 52'N 39°53T, 610 m o,d., 1.-2. XI. 1986, W.BUttiker. Saudi
Arabia: Tabuk, Wadi Sadr, 27" 53'N 3S o 46'E, 620 m o.d., 5.-7. XI. 1986, W, Btittiker. - 2 dd, Saudi Arabia: Makkah, Wadi
Shuqub, 20° 43'N 41 0 10'E, 1390 m o.d., 6. IV. 1980, W. Btittiker. - 2 dd, Saudi Arabia: Makkah, Wadi Turabah, 20 0 29'N
41°15'£.,1400 m o.d., \\I,Buuiker. - 3 dd, Saudi Arabia, Riyadh, Wadi Sheib Luha, 24°27'l'i 46°12'E, 575 m o.d"
14.-18.VIII.1978, W.Btittiker. - 5 dd, Saudi Arabia: Eastern Province, Hofuf, 25 c 24'N 49°28'E, 150 m o.d., 28,V.1978, \\I.
Buttiker (2 to NHMB). - 2 dd, Saudi Arabia: Medina, Fare, 1160 m o,d., 23°05'N 39°54'E, 13.-14.XI.1986, W, Buttiker. - 1
d, Saudi Arabia: Baha, \Xfadi Ibrahim, 20° 24'N 41 ° 11T" 1790 m o.d., 24,-26. VIII. 1985, W. Btittiker. - I d, Saudi Arabia,
Baha, Al Mindak, 20"07'N 41 ° IS'E, 1760 m o.d., 8. IV. 1980, W. Bilttiker. - 1 d, Saudi Arabia: Asir, 18° 14'N 42° 56'E, 2250 m
o.d., IS. VII. 1981, \\I. BUttiker. - 5 dd, Saudi Arabia: Najran, Talaa, 18°04'I\: 43° 57'E, 1420 m o.d., W. Btittiker.
Hypopygium: fig. 12.
Distribution: Palaearctic region: Algeria. A specimen from Jordan ('Palestine, Jordan valley',
Jericho, 18. IV. 1923, P.A. Buxton, BMNH) is a first record for Jordan. Afrotropical region: Zimbabwe,
South Africa.
,

FAUNA OF SAUDI ARABIA 10, 1989
Comments: Paramerina vailfanti is one of the most widely distributed species of Saudi Arabian
Chironomidae, occurring from Al Jawf in the north to Talaa in the south and at altitudes ranging
from 150 m at Hofuf to 2260 m at Abha. This extensive material shows greater variation than has
been recognised previously, particularly in the antennal ratio which can vary in a single population
Qawf) from 1.28-1.5, and reaches a maximal value of 1.65 in a specimen from Fare. As a generalisation,
specimens from higher altitudes are larger and have a higher antennal ratio. Saudi Arabian specimens
blur the distinction between P. mal1retanica Fittkau and P. vail/anti, but synonymy must await
examination of the immature stages, particularly the pupae, of a range of Saudi Arabian specimens.
Paramerina sp. 1 sensu LEHMA:-lN (1979)
Paramerina sp. I Lehmann, 1979. - Spixiana Supp!. J: 18 (part of original material examined, 2SM).
Ma terial: 1 cJ, Saudi Arabia: .\1edina, Fare, 23 °05'N 39° 54'E, 1160 m o.d., 13.-14. XI. 1986, W. Bilttiker. - 1 cJ, Saudi
Arabia: Makkah, Wadi Maraum, 22° 16'N 39°44'£, 280 m o.d., 3.-4. V.1984, W. BUttiker. - 1 cJ, Saudi Arabia: Baha,Wadi
Diyhan, 19" 4S'N 41 0 36'E, 830 m o.d., 7. III. 1984, W. Bilttiker.
Comments: The three males above differ from P. vail/anti in genitalic, particularly paramere,
structure, spur conformation and the distinctly more elongate palp segments. The closest species
appears to be LEHMANN'S (1979) unnamed Paramerina from Zaire, but confirmation of identity with
the variably damaged Saudi specimens cannot be made. The material is too damaged for illustration,
but the hypopygium seems to closely resemble LEHMANN'S (1979) fig. 28.
Subfam. Orthocladiinae
Key to the species of Saudi Arabian Orthocladiinae
1. Eye hairy, with macrotrichia projecting beyond ommatidia. Tergite IX lacking anal
point .......................................... . 2
Eye bare. Tergite IX with distinct anal point .................... . 3
2. Scutum with dorsocentral setae erect, arising from distinct, often pale, pits. Abdomen
more or less unicolourous, dark Paratrichoc1adius micans
Dorsocentrals decumbent, not arising from distinct pits. Abdominal tergites I and IV
pale, remainder dark Cricotopus (Cricotopus) albitibia
3. Wing membrane with dense macrotrichia Paraphaenocladius impensus
Wing membrane bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... . 4
4 Pulvilli absent. Hypopygial penis sac with very strong virga
Bryophaenocladius paraproductus
Pulvilli very broad, pad-like, subequal in length to claw. Virga absent
Psectrocladius limbate//us
Bryophaenocladius paraproductus n. sp.
Material: Holotype: cJ, Saudi Arabia: Makkah, Harithi. 21 0
l8'N 40° 18'E, 1910 m o.d., 23.-24. V.1985, W. BUttiker
(BMNH).
Des crip ti on: Body length 2.25 mm. Body colour bleached, but apparently brown, vittae scarcely
darker.
Antennae missing; eye with slight dorsomedial projection, with 6-7 verticals, without postorbitals;
palp segment II-V lengths: 43, 87, 87, 130 J,Lm, lacking any sensilla on segment III.
Acrostichals not visible on damaged thorax; 4-5 fine lateral antepronotals, 11-12 uniserial dorsocentrals,
4 prealars, 10 uniserial scutellars.
245

246 P. S. CRANSTON, D. D. JUDD
Wing length 1.2 mm, costal extension about 60 11m beyond apex of R4+5' venarum ratio 1.35, CUt
curved, but not sinuous. Vein R with 2 setae, remaining veins apparently bare.
Anterior leg ratio 0.65.
Hypopygium as in fig. 13.
Distribution: Bryophaenocfadiu5 paraproductus is known only from the holotype locality, Harithi,
in Makkah, Saudi Arabia.
Co mme n ts: Bryophaenocfadiu5 paraproductus completely conforms to the generic diagnosis given
by CRANSTON et al. (1989), belonging to the majority group of species lacking a digitiform extension
to palp segment 3 and with a fully developed hind tibial comb, although with a reduced number of
4-5 spines therein.
Without a cladistic analysis of the genus it is not possible to determine the relationships of this
species, but the Afrotropical species Bryophaenocfadius productu5 (Freeman) bears a close resemblance
on non-genitalic features and has a somewhat similar hypopygium. As the specific epithet implies, B.
paraproductus might be sister species to B. productus.
Cricotopus (Cricotopus) albitibia (Walker, 1848)
Chironomus albitibiaWalker, 1848. List of the specimens of dipterous insects in the collection of the British Museum 1:
16. Type-locality: Sierra Leone (Holotype d, pinned, examined, BMNH).
Cricotopus albitibia. - Freeman 1956; Bull. Br. Mus. ;..rat. Hist. Ent. 4: 306.
Material: 2 dd, Saudi Arabia: Gizan, \Yadi Damad, 17° 17'N 43"06'E, 800 m o.d., 24. IX. 1981, W. BUttiker.
Hypopygium: fig. 14.
Distribution: Widespread in Afrotropical region, from Ethiopia and Sudan southwards. The
present finding is the first record from outwith the Afrotropical region.
Comments: The distinction between the Afrotropical C. albitibia and the Holarctic C. bicinctu5
is uncertain in the adult stage. We accept FREEJ\1l\N'S (1956: 306) observation of hypopygiaJ differences
that are evident in Arabian material, but have reservations that will only be resolved by the discovery
of the immature stages in Africa.
Paraphaenocladius impensus (Walker, 1856)
Chironomus impen5lls Walker, 1856. - Ins. Brit. Dipt. 3: 184. Type-locality: England (Holotype examined, BMI\H).
Paraphaenocladius impensus. Pinder 1978; Scient. Rep. Freshwat. BioI. Ass. 37: 92.
Mate ri al: 2 dc!, Saudi Arabia: '\1akkah, Khoda, 19.IX. 1982,20° 2tN 41 0 1 S'E, 1890 m o.d., W. Btittiker. - 1 Saudi
Arabia: Baha, Wadi Diyhan, 830 m o.d., 19° 48'N 41 0 36T, 7. III. 1984, W. BUttiker.
Hypopygium: fig. 15.
Distribution: widespread in western Palaearctic including Lebanon and Algeria; Canada. Specimens
from Turkey (1 d, Konya, 20 miles [32 km] SW of Konya, 32°24'E 3r4tK, swept beside
stream, 4,000' [1220 m o.d.], 13. VIII. 1974, P. S. Cranston, BM 1974-482. - 1 d, Tatvan, 3 km S Ahlat,
swept from reedy vegetation beside wet bog, 19.\'111.1974, 5500' [1677 m o.d.], 19.VIII.1974, P.S.
Cranston, BM 1974-482 are the first records of the species from Turkey.
Paratrichocladius micans (Kieffer, 1918)
Dactylocladiu5 micans Kieffer, 1918b. - Ann. hist.-nat. Mus. nat. Hung. 17: 81. Type-locality: South Africa, Natal [Type(s)
not examined].
TrichocladiH5 micans. Freeman 1956; Bull. Br. Mus. Nat. Hist. Ent. 4: 314.
Material: 5 Saudi Arabia: Makkah, Wadi Nimar, 21 °OS'N 40° 58'E, 1500 m o.d., 19.-20. V. 1983, W. Biittiker (I to
NHMB, 1 to ZSM). 1 d, Saudi Arabia, Asir, Abha, 18 c 14'N 42 0
56'E, 2260 m o.d., 15. VIII. 1981, W. Btittiker. 1 d, Saudi
Arabia, Asir, Wadi Ash Sharayi, 18 c 04'N 43 0
38'£, 24.-25. IX. 1979, W. BOttiker .
Distribution: Palaearctic region: Greece, Lebanon. Afrotropical region: Yemen, Ethiopia and
eastern subSaharan Africa to South Africa, Madagascar and Reunion.

14
FAUNA OF SAUDI ARABIA 10, 1989
1
I
t!1\ )
I
\
I
!
Figs 14-17: 14, Cricatopus Cricatopus) albitibia, hypopygiunL 15, Paraphaenodadim impensus, hypopygium, 16, Parat1"ichocladius
micans, hypopygium. 17, PsectrocladiT;s limbatellus, hypopygiulll.
Four males from Turkey (Antalya, Tara beach, 26. v'lIl. 1974, P. S. Cranston, BM 1974-482, from
beside the splash zone of waterfall) are the first record of the species from Turkey. REISS (1985) records
the congeners P. rufiventris from far eastern Turkey, and P. skirwithensis from the Turkish eastern
Black Sea coast.
Psectrocladius limbatellus (Holmgren, 1869)
Chironomus limbatellus Holmgren, 1869. - K. Svenska Vet.-Akad. Hand!. [4 J8(5): 44. Type-locality: Spitzbergen (Lecto­
type examined 1982, SNHM)
Psectrodadius timbatelhls. - Langton, 1984; Ent. Seand. 15: 479 (nee Psectrocladius limbatellus. - auctt. ante 1984).
Psectrocladius edward5i Brundin, 1949. - Rep. Inst. I'reshwat. Res. Drottninghollll 30: 816.
Psectrodadius edwardsi. Langron, 1980; Ent. Gaz. 31: 79.
Material: 2 cfcf, Saudi Arabia: Jaw£. AlJawf, 29° 52'N 39° 53'£,610 III o.d., 1.-2. X!. 1986, W. BUttiker. - 2 Saudi
Arabia: Riyadh, Al Ghat, 26°07'N 44°58'E, 730 m o.d., 17.-18.X.1978, W.Btittiker.
247

248
P. S. CRAl"STON, D. D. JUDD
Hypopygium: fig. 17.
Comments/Distribution: Psectrocladius limbatellus, as recognised by re-examination of the
type series, is the species previously referred to as Psectrocladius edwardsi Brundin. LANGTON (1980)
viewed this as an essentially Palaearctic boreal species, but from his subsequent paper (LANGTON 1984)
it is clear that the broader species concept includes at least eurythermic southern English populations.
Given the previous identificatory difficulties, and the probable requirement of pupal exuviae to confirm
identity, the true distribution is unclear.
A specimen from Turkey (Tatvan, Adilcevac, slopes of Suphan Dagi, 6600' (2015 m o.d.],
19. VIII. 1974, p.s.Cranston, BM 1974-482) is virtually identical to Saudi Arabian specimens, differing
only in possessing a rather blunt apex to the gonostylus in contrast to the usual rounded curve.
These constitute first records of the genus Psectrocladius from Turkey and Saudi Arabia. In the
Afrotropical region the genus is known only from a single species from South Africa.
Subfam. Chironominae
Key to males of Saudi Arabian Chironominae
1. Wing membrane bare, or, when with macrotrichia, squama fringed. Chironomini 2
Wing membrane with macrotrichia, or, when bare, as with all other members of the
tribe, lacking squamal fringe T anytarsini 26
2. Antenna with 11 flagellomeres. Hind tibial combs strong, separate and each bearing
strong spur. Pulvilli well developed, pad-like, and subequal in length to claw . . . . ..
Antenna with 13 flagellomeres. Hind tibial combs may be reduced, more or less fused
and often lacking one or sometimes both spurs. Pulvilli poorly developed, usually
3
3.
shorter than claw length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14
Inferior volsella (i. v., figs 19, 22) a distinct, densely microtrichiose and setose lobe,
extending posteriorly at least to level of apex of anal point . . . . . . . . . . . . . . .. 4
4
Inferior volsella absent or very small; never an elongate setose lobe ..... . . . . .. 10
Inferior volsella narrowest medially, bowed in a dorsoventral plane, broadened to bifid
apically ..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
Inferior volsella more or less clubbed, never narrowest medially and neither bowed nor
5
bifid apically ......................................... 7
5. Inferior volsella apically broadened but not bifid (fig. 25). Posterior contour of tergite
IX without hyaline appendage Dicrotendipes binotatus
Inferior volsella apically bifid (fig. 26). Hyaline appendage postero-Iateral on tergite
IX, lateral to anal point base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6
6. Dark mark medially in wing cell m1+2 (fig. 23) Dicrotendipes peringuey:U1us
Wing cell m1 + 2 without medial dark mark Dicrotendipes sudanicus
7. Superior volsella elongate ovoid with setae and microtrichia on dorsomedial surface,
bare laterally; without digitiform process unknown genus
Superior volsella with setose/macrotrichiose pad-like base and variably digitiform bare,
medially directed, extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8
8. Pad-like base of superior volsella large, extending posterior to digitiform part, which
arises dorsally from the baso-Iateral corner of the pad Kiefferulus disparilis
Pad-like base of superior volsella smail, lying posterior to, and in virtually same plane
as, digitiform extension Cbironomus 9
,

fAUNA OF SAUDI ARABIA 10,1989
9. Femoral apex darkened, separated from dark tibial apex by pale "knee". Wing markings
restricted to darkened cross vein MCu Chironomus seyche11eanus
Legs essentially unicolourous. Wing markings patterned as fig. 18.
Chironomus caJipterus
10, Inner margin of gonocoxite with two appendages the superior volsella more or less
rounded and partially or completely covering the inferior volsella (figs 21, 22), Lobes
of tergite IX absent Cryptochironomus 11
Inferior volsella absent, superior volsella elongate. Setose lobes lying lateral to anal
point, ventrally on anterior margin of tergite IX (figs 36-38) Microchironomus 12
11. Anal point somewhat bulbous apically. Volsellae not widely disparate in size, inferior
volsella visible lateral to superior volsella (fig. 21) Cryptochironomus diceras
Anal point tapering to rounded point. Volsellae disparate in size, the larger superior
volsella obscuring the inferior (fig. 22) Cryptochironomus rostratus
12. Anal point apically broadened to rounded apex. Gonostylus broadest at base. Superior
volsella short (fig. 36) Microchironomus deribae
Anal point parallel-sided or tapering to blunt point. Gonostylus either of equally wide
through its length or broadest submedially. Superior volsella elongate , . . . . . . . .. 13
13. Anal point tapering to point, mediobasally with ventral setose pad. Gonostylus broadened
submedially (fig. 37) Microchironomus lendJi
Anal point parallel-sided, with ventral setose pad at base, Gonostylus of more or less
even width (fig. 38) Microchironomus tener
14. Median volsella (m.v" fig. 42) present as a distinct setose lobe. Squama bare
Paratendipes 15
Median volsella absent, represented, at most, by a wart bearing few simple setae 16
15. Wing with bold pattern (fig. 41) Paratendipes nubiJipennis
\Ving with only faint darkening around anterior veins Paratendipes nudisquama
16, Scutum with strong tubercle (fig. 51) Stictochironomus puripennis
Scutum lacking tubercle ... , ......... , .. , ........... , . . . . .. 17
17. Inferior volsella lacking strong elongate apical seta (fig. 39). Trace of median volsella
present. Fore tibial apex without scale or spur Microtendipes umbrosus
Inferior volsella always with strong, elongate, posteriorly directed apical seta. No evidence
of median volsella. Fore tibia with scale, usually either developed as, or bearing
subapically, a spur (e.g. figs 48, 50) Polypedilum 18
18, Wing membrane with dense macrotrichia. Hypopygium as in fig. 43
Polypedilum (Pentapedilum) wittei
Wing membrane bare . . . , . . , . , , . . . , . . , . , . . . . . . , . . . . , . . . . ,. 19
19. Anal point with small lateral notches. Base of anal point, on tergite IX, with variably
developed lateral projections (fig. 49). Gonostylus very narrow
Polypedilum (Tripodura) aegyptium
Anal point smooth, with neither lateral notches nor basal projections, Gonostylus not
abnormally narrow . . . . . . . . . . . . . . . , . . . . . . . . 20
20. Wing with dark markings (maybe faint in bleached specimens) . . . . . . . . . . 21
Wing unmarked .. , .. , ... , . , ..... , ... , ..... , . . . . . . . . 25
21. Gonostylus with characteristic preapical contraction, then apical bulge (fig. 44)
Polypedilum (Polypedilum) alticola
Outer margin of gonostylus evenly curved to apex . . . . . . . . . . . . , . . . . . . .. 22
249

FAUNA OF SAUDI ARABIA 10, 1989 251
34. Anal point without groups of spines. . . . . . . . . . . . . . . . . . . . . . 35
Anal point dorsally bearing groups of spines . . . . . . . . . . . . . . . . . 36
35. Digitus (appendage la) absent. Anal point somewhat lanceolate (fig. 60)
Tanytarsus pallidulus
Digitus present. Anal point parallel sided Tanytarsus nimar
36. Median volsella short, comprising three broad lobes (fig. 62). Digitus characteristically
curved, almost corkscrew-like, extending well beyond median margin of superior volsella
(fig. 61) Tanytarsus trifidus
Median volsella longer, with lanceolate setae. Digitus short, not extending beyond
median margin of superior volsella ....... . . . . . . . . . . . . . . . . . . . . .. 37
37. Oro-lateral margin of superior volsella with group of macrotrichia (fig. 58)
Tanytarsus horni(l)
Oro-lateral margin of superior volsella without macrotrichia (fig. 57)
Tanytarsus mcmillani(1)
Chironomus calipterus Kieffer, 1908
Tribe Chironomini
Chironomus calipterus Kieffer, 1908. Denkschr. med.-naturw. Ges. Jena. 13: 158. Type-locality: S.W. Africa (',,'amibia),
Rooibank [Type(s) not examined].
Chironomlls calipterus. - Freeman 1957; Bull. Br. Mus. ',,'at. Hist. Ent. 5: 345.
Material: 2 0'0', Saudi Arabia, Jawf, Al Jawf, 29° 52'N 39° 53'E, 610 In o.d., 1.-2. Xl.! 986, W. BUttiker. Saudi
Arabia: Makkah, Wadi Hanaq, 22° 44'N 39° 22'E, 100 m o.d., W. BUttiker. 2 0'0', Saudi Arabia: Eastern Province, Hofuf,
25°24'N 49°28'E, 150 m o.d., W.Blittiker. - 6 Saudi Arabia, Al Hassa Oasis, canal, 26.XI.1981, W.BUttiker. - 4 0'0',
Oman: Tahwa nr Sur, 'light trap in goat pen' 25.XL 1986, R. P. Lane (I to NH.'vlB). 20'0', larvae. United Arab Emirates,
Zakhar Alain, 27. IV. 1980, G. B. White.
Wing: fig. 18.
Hypopygium: fig. 19.
Distribution: Palaearctic region: circum-Mediterranean from southern France to Palestine,
Syria. A specimen from Jordan (Azraq Shishan marsh, 21.IY.-ll.V.I966, D.S. Fletcher, International
Jordan Expedition, BM 1966-298) is a first record for the country. Widespread in Afrotropical
regIOn.
Chironomus seychel1eanus Kieffer, 1911
Chironomlls seychelleanlls Kieffer, 1911h. Trans. Linn. Soc. Lond. (2. Zool.) 14: 356. Type-locality: Seychelles: Mahe
(Lectotype 0' examined, BMl'\H).
Chironomlls callichirus Kieffer, 1911b. Trans. Linn. Soc. Lond. (2. Zool.) 14: 352 (preoccupied by ChironomHs ca{fichims
Kieffer. 1911a. Type-locality: Seychelles).
ChironomHs mahensis Kieffer. 1912. - Boll. Lab. ZooI. gen. Agr. Portiei 6: 175 [replacement name for Chironomus caflichirus
(Kieffer), preoccupied Kieffer, 1911 a: Rec. Indian Mus. 6:
Material: 20'0', Saudi t\rabia, Jawf, Al Jawf, 29°52'N 39°53'E, 610 m o.d., 1.-2. XI. 1986, W.BUttiker (1 to ZSM). - 10',
Saudi Arabia: Makkah, Harithi, 21 0
18'N 40" 18'E, 1910 m o.d., 23.-24. V. 1985, W. BUttiker. - 1 0', Saudi Arabia: Makkah,
Wadi Mirtad, 20" 53'N 40° S3'E, 1500 m o.d., 31. VIII. 1984, W. BUttiker. - 1 0', Saudi Arabia: Asir, As Sudr, 18 ° OO'N 42° 38'E,
1220 m o.d., 8. IX. 1984, W. Biittiker. 1 Saudi Arabia: Asir, Rabifa, 18°01'N 42° 46'E, 2320 m o.d., 7. IX. 1984, W. Biittiker.
- 1 0', Saudi A£abia: Najran, Talaa, 18 "04'N 43 0
S7'E, 1420 m o.d., W. Blittiker. 10', Oman: Tahwa nr Sur, 'light trap in goat
pen' 25.XI. 1986, R.P.Lane.
(1) See comments under these species in following text.

252 P. S. CRANSTON, D. D. JUDD
Hypopygium: fig. 20.
Distribution: Afrotropical region: widespread eastern and southern Africa, including Seychelles,
Madagascar and Reunion. This constitutes the first record from outside the Afrotropical
reglOn.
Com ment s: C. seychelleanus is amongst the more distinctive species of Chironomus in possessing
dark apices to the femora separated by a pale 'knee' from the darkened base of the tibia. Furthermore,
the cross vein (wing vein MCu) is darkened, as is a central area of the anterior tergites.
Cryptochironomus diceras Kieffer, 1923
Cryptochironomus diceras Kieffer, 1923. - Ann. Soc. ent. Fr. 92: 163. Type-locality: Sudan: Shamhe [Type(s) not
examined].
Cryptochironomus diceras. - Freeman 1957; BulL Br. Mus. Nat. Hist. Ent. 5: 390.
Material: I Saudi Arahia: Eastern Province, Hofuf, 2So24'N 49°28'E, 150 m o.d., 26.VL 1978, W.Btittiker. - I
Saudi Arahia: Eastern Province, Hofuf, 25°24'N 49°28'E, 150 m o.d., IS.XL 1985, W.Peters, BM 1986-262.
Hypopygium: fig. 21
Distribution: widespread in the Afrotropical region: from Sudan and Ethiopia southwest and
southwards to South Africa. This constitutes the first record from outside the Afrotropical region.
Comments: Species delimitation in CryptochironomtJS is unclear and relies to a large extent on
somewhat variable colour patterns, lacking detailed assessment of genitalic structure. However, C.
diceras appears distinctive by virtue of the stout anal point and shape of the volsellae. The two Saudi
Arabian specimens of C. diceras have a more bulbous apex to the anal point than is illustrated by
FREEMAN (1957: fig. lIe), but examination of further material from Africa shows that the present
material is not atypical.
Cryptochironomus rostraWs Kieffer, 1921
Cryptochironomus rostratus Kieffer, 1921 a. - Bull. Soc. Hist. nat. Metz 29: 67. Type-locality: (Poland) Silesia [Type(s) not
examined].
Cryptochironomus rostratu5. - Pinder 1978; Scient. Rep. Freshwat. BioI. Ass. 37: 116.
Material: 1 Saudi Arabia: Abah, 4. VII. 1983, A.S.Talhouk (no further details provided).
Hypopygium: fig. 22.
Distribution: Palearctic region, from Britain eastwards to Lebanon, Turkey and Syria.
Comments: C. rostratus is distinctive within the genus Cryptochironomus by virtue of the small
(no longer than wide) frontal tubercles, the parallel-sided anal point and the virtually round superior
volsella.
A specimen from NE Syria (Khabur, Tall Shaikh Hamad, 35 37'N 40045'E, 20.X.-9.XI.1986,
F. Krupp & W. Schneider) adds to REISS' (1986) records from northwestern Syria.
Dicrotendipes freemani Epler, 1988
Dicrotendipes /reemani Epler, 1985. Mem. Am. Ent. Soc. 36: 37. Replacement name for Chironomus binotatus Kieffer,
1911 h preoccupied hy Chironomus binotatHs Wiedemann, 1817.
Chimnomu5 binotatus Kieffer, 1911b. Trans. Linn. Soc. Lon. 14: 356. Type-locality: (Seychelles) Mahe (Holotype 9
examined, BMNH).
)Chironomus (Limnochironomus) chambiensis Goetghebuer, 1936. Revue ZooL Bot. Afr. 28: 464. Type-locality: Zaire:
Kabasha, Chambi [Type(s) not examined].
Chironomus (Dicrotendipes) binotatus. - Freeman 1957; Bull. Bf. ",1us. Nat. Hist. EnL 5: 367.
Materia1: 1 Saudi Arabia: Eastern Province, Hofuf, IS.XI. 1985, W. Peters, BM 1986-262 (BMNH). - 4 ad, Oman:
Tahwa nr Sur, 25.XL 1986, R.P.Lane (1 to ZSM, 3 to BMNH).
Hypopygium: fig. 25.
Distribution: Afrotropical region: Seychelles, Ethiopia (EPLER, 1988). These specimens from
Saudi Arabia and Oman constitute the first records from outside the Afrotropical region.

FAUNA OF SAUDI ARABIA 10, 19,9
22
" I
'/
5)! \
U
Figs 18-22: 18,19, Chironomus calip/ems, 18, wing, 19, hypopygium. 20, Chironomus seychelleanus, hypopygium (right side, dor­
sal, only). 21, CryptochiroHomus diceras. hypopygium. 22, Cryptochironomus rostratus, hypopygium.
Comments: As noted by EPLER (1988), D.freemani is extremely similar to D. chambiensis and differences
noted by FREEMA:-I (1957) (as Chironomus (D.) binotatus) include artefacts of genitalic slide
preparation. EPLER (1988) concludes that synonymy between D. freemani and D. chambiensis cannot be
made without knowledge of the immature stages. We chose to apply the name D. freemani because
253

254 P. S. CRANSTON, D. D. JUDD
specimens so identified in BMNH have a narrower base to the anal point and more setae on tergite
IX than do any identified D. chambiensis. A difference of uncertain significance between Arabian D.
binotatus and Afrotropical material in the BMNH concerns the number of setae towards the hyaline
apex of the superior volsella - Arabian material has three, of which one is stouter - all previously collected
specimens have four subequal finer setae.
Dicrotendipes peringueyanus Kieffer, 1924
Dicrotendipes peringueyanus Kieffer, 1924. Ann. Soc. scient. Brux. 43: 257. Type-locality: South Africa; Cape Province
[Type(s) not examined].
Chironomus (Dicrotendipes) peringueyanus. - Freeman, 1957: Bull. Br. Mus. Kat. Hist. Ent. 5: 365.
Dicrotendipes peringueyanus. - Contreras-Lichtenberg 1986; Ann. naturh. Mus. Wien 88/89: 706.
Dicrotendipes peringueyanus. Epler 1988; .Mem. Am. Ent. Soc. 36: 41.
Material: 1 d, Saudi Arabia: Medina, Khaybar, 680 m o.d., 25° 36'K, 39° E, 26. IV. 1979, W. BUttiker. - Saudi
Arabia: Baha, Adnan, 1350 m o.d., 20° 26'K 41 0 3 I'E, 21.-23. IX. 1978, W.BUttiker.
Wing: fig. 23.
Dis tribution: Palaearctic Region: France, Spain, Algeria; Afrotropical regIOn: Chad, Kenya,
Senegal, Zaire, Zambia, Zimbabwe.
Figs 23-26: 23, Dicrotendipes peringueyanus, wing. 24, Dicrotendipes sudanicus, wing. 25, Dicrotendipes /reemani, hypopygium. 26,
Dicrotendipes sudanicus, hypopygium.
23
24

FAUNA Or SAUDI ARABIA 10, 1989
27-35: Kieffirul14s disparilis. 27, larval mentum. 28, dorsal surface of larval head. 29, larval mandible. 30, larval antenna. 31,
larval labrum-epipharynx, 32, basal ring of pupal thoracic horn. 33, apico-lateral corner of pupal tergite IX. 34, dorsal tergites
of pupa. 35, hypopygium.
257

258 P S. CRANSTON, D. D. JUDD
Microchironomus JendH (Kieffer, 1918)
Chironomus (MicrochironomtlS) lendli Kieffer, 1918b. - Ann. hist.-nat. Mus. nat. Hung. 16: 112. Type-locality: (Turkey)
'Asie-Mineure, Afium-Karahissar' not examined].
Chironomomus (Cryptochironomus) Freeman,1954. Proc. Roy. Ent. Soc. Lond. (B) 23: 176. Type-locality: (South
Africa) Berg River, Piquetberg o examined, BMNH].
ChironomomtlS (Cryptochironomus) Freeman 1957; Bull. Br. Mus. Nat. Hist. Ent. 5: 395.
Material: 1 Saudi Arabia: Gizan, Ad Darb, 1 r 44'N 42° 14'E, 64 m o.d., 9.-10. III. 1979, W. Blittiker.
Hypopygium: fig. 37.
Distribution: Palaearctic region: Turkey, Palestine and Lebanon. Widespread in Afrotropical
region from Sudan southwards to South Africa.
Microchironomus tener (Kieffer, 1918)
Chironomomus (Cryptochironomus) tenerKieffer, 1918a. Ent. Mitt. 7: 48. Type-locality: (Poland) Lazarettschiffzug Dan­
zig [Type(s) no longer extant, not examined].
ChironomomJ,ls (Cryptochironomlls) forcipatus Freeman, 1954. PJ'oc. Roy. Ent. Soc. Lond. (B) 23: 20. Type-locality:
(South Africa) Berg River, Piquetberg (Holotype 0 examined, BMNH).
Chironomomus (Cryptochironomlls) forcipatus. Freeman 1957; Bull. Br. Mus. Nat. Hist. Ent. 5: 394.
Mate rial: 1 0, Saudi Arabia: Gizan, Ad Darb, 1 r 44'N 42° 14'E, 64 m o.d., 9.-10. III. 1979, W. BUttjker.
Hypopygium: fig. 38.
Distribution: Palaearctic region: widespread from Britain eastwards to the Soviet far East and
North Korea, southwards to Egypt and Palestine. Afrotropical region: widespread, south to South
Africa. Oriental region: India to Indonesia. Austropacific region: Australia.
Microtendipes umbrosus Freeman, 1955
Microtendipes umbrosus Freeman, 1955a. Explor. Pare )jato Albert Miss. Witte 83: 32. Type-locality: Kenya, Nyanza,
Lumbwa District (Holotype y, paratype 0 examined, BMNH).
Microtendipes IImbro5Us. - Freeman 1958: Bull. Br. Mus. Nat. Hist. Ent. 6: :'113.
Material: 2 0d', Saudi Arabia: Makkah, Harithi, 21 0 18'N 40° 1sT, 1910 m o.d., S.X. 1984, W. BUttiker.
Wing: fig. 40.
Hypopygium: fig. 39.
Distribution: Afrotropical regIOn: Southern Africa and at increasing altitude northwards to
Nigeria, Sudan and Ethiopia.
Paratendipes nubiJipennis Freeman, 1957
Paralendipes nllbilipennis Freeman, 1957. - Bull. Br. Mus. Nat. Hist. Ent. 5: 422. Type-locality: N. Nigeria, Alixaga
(Holotype 0 examined, BMNH).
Material: I 0, Saudi Arabia: Makkah, Wadi Mirtad, 20° 53'N 40° 53'E, 1500 m o.d., 31. VIII. 1984, W. BUttiker. 1 0,
Saudi Arabia: Wadi Shuqqub, 20° 43'N 41 0
10'E, 1380 m o.d., 12. IX. 1986, W. BUttiker. 2 Saudi Arahia: Baha, Adnan,
20026'N 41°31'E, 1350 m o.d., 21.-23.IX.1978, W.BOttiker (l NHMB, 12SM). 10, Saudi Arabia: Baha, Wadi Ibrahim,
20° 24'N 41 ° 11 T, 1790 m D.d., 24.-26. VIII. 1985, W. Biiuiker. - 1 0, Saudi Arabia: Asir, Khamis Mushayt, 18° 19'N 42°43'E,
1890 m o.d., 29. IX. 1978, W.Btittiker.
Wing: fig. 41.
Hypopygium: fig. 42.
Distribution: Afrotropical region: Nigeria, Burkino Faso (Upper Volta), Zambia, Zimbabwe.
Comments: The wing pattern, distinctive in the holotype and some other dry pinned museum
specimens, is less clear in Saudi Arabian material, although visible under transmitted light and with
phase contrast. Either bleaching has occurred or these northern populations are paler.
Paratendipes nudisquama (Edwards, 1929)
Chironomus (Paratendipes) nudisquama Edwards, 1929. - Trans. Roy. Ent. Soc. Lond. 77: 396. Type-localities: &
Ireland (Syn types 00 examined, BMNH).

[
FAUNA OF SAUDI ARABIA Ie, 1989
Figs 36-39: 36, Microchironomu5 deribae, hypopygium. 37, Microchironomu5 lendli, hypopygium. 38, Microchironomus tener,
hypopygium. 39, Microtendipes umbroSlJs, hypopygium.
37
259

260 P. S. CRANSTON, D. D. JUDD
Material: 2 d'd', Saudi Arabia: Gasim, Khubra Barn, 26°05'N 43° 39'E, 15.-16.X.1978, W.Blittiker. - Several d'd' (in
alcohol), Saudi Arabia: Makkah, Wadi Ellah, 20° 35'N 41 0
36'E, 1480 m o.d., 9. IX. 1983, W. Bilttiker. Several d'd' (in alco­
hol), Saudi Arabia: Makkah, Wadi Hanaq, 22° 44'N 39°22'E, 100 m o.d., W.Blittiker. 7 d'd', Saudi Arabia: Makkah, Wadi
lvlaraum, 2r22'N 39° 44'E, 280 m o.d., 13. V. 1984, W. Blittiker. - I Saudi Arabia: Makkah, Wadi Mirtad, 20° 53'N 40° 53'E,
1500 m o.d., 31. VIII. 1984, W. Bilttiker. - I d', Saudi Arabia: Makkah, Wadi Shuqub, 20° 43'N 41 0 10'E, 1390 m o.d., 6. IV. 1980,
W. Bilttiker. - 1 d', Saudi Arabia: Makkah, Wadi Turabah, 20 0 29'N 41 0 15'E, 1400 m o.d., W. BUttiker. - 1 d', Saudi Arabia:
Eastern Province, Hofuf, 25 ° 24'N 49° 28'E, 150 m o.d., 28. V. 1978, W. BUttiker. - 9 d'd', Saudi Arabia: Baha, Adnan, 20° 26'N
41°3rE, 1350 m o.d., 21.-23. IX. 1978, W.Blittiker (I NHMB, 1 ZSM). 1 d', Saudi Arabia: Baha, Wadi Diyhan, 19°48'N
41°36'E, 1350 m o.d., 7. II1.I984, W. BUttiker. 2 d'd', Saudi Arabia: Baha, Wadi Ilyab, 20° 24'N 41 0 JO'E, 3. II. 1984, 160 m
o.d., W. BUttiker. - 2 d'd', Saudi Arabia: Asir, Wadi Ash Sharayi, 18 0 04'N 43 0 38'E, 24.-25. IX. 1978, W. BUttiker. - I d', Saudi
Arabia: Gizan, Hakimah, 17"02'N 42°57'£:, 14.-16.X.1979, 100 m o.d., W.BUttiker. I d', Saudi Arahia: Gizan, Maraba,
IrS4'N 42° 23'E, 320 m o.d., l.X.1978, W.BUttiker. - 3 d'd', Oman: Tahwa nr Sur, 'light trap in goat pen', 25.XI.1986, R.I'.
Lane.
Distribution: Palaearctic region: widespread, from Scandinavia and Britain eastwards to
Greece, Lebanon and Transcaucasus.
Comments: The genus Paratendipes is in need of revisionary study. Two species groups are easily
distinguished: the P. albimanus group has squamal setae and a slightly broadened anal point, while
the P. nudisquama group has a bare squama and tapering anal point. Within the P. nudisquama group
the genitalia are remarkably uniform throughout the described species; major specific differences are
found in the wing and leg pattern and colouration. P. nudisquama is recognised in this study by the
absence of squamal setae, uniformly mid-brown coloured legs and lack of a wing pattern. In the Afrotropical
region the widespread P. crosskeyi Freeman lacks squamal setae and wing pattern, but has all
femora and the fore tibia darkened apically; the hypopygium is indistinguishable from P. nudisquama.
Another Afrotropical species, P. striatus Kieffer, with a faint wing pattern that might represent a
darker version of the wing vein infuscation seen in some P. nudisquama, also has some banding on the
legs, and but has an anal point with a somewhat bulbous apex. We conclude that the Arabian specimens
recognised above, lacking a distinct wing and leg pattern, all represent P. nudisquama, despite
the apparent wide ecological range of habitats from which it has been taken.
Polypedilum (Pentapedilum) wittei Freeman, 1955
Polypedilum wittei Freeman, 1955. Explor. Pare. Nat. Albert Miss. Witte 83: 30. Type-locality: (Zaire) Kivu, Kalondo
[Type(s) not examined].
PolypediIum wittei. Freeman 1958; Bull. Br. Mus. Nat. Hist. Ent. 6: 301.
Material: 1 d', Saudi Arabia: Baha,AI Foqa, 19°50'N 41"15'E, 1630 m o.d., 10.IX.1984, W.BUttiker.
Hypopygium: fig. 43.
Distribution: Afrotropical region: Ethiopia, Sudan, Chad and Guinea, Nigeria south to Transvaal.
This constitutes the first record from outside the Afrotropical region.
Comments: P. (Pentapedilum) wittei bears a rather close resemblance to the Palaearctic P. (Pentapedilum)
uncinatum Goetghebuer 1921, differing in the much paler general body colour, and the presence
of narrow dark bands at the apices of the abdominal segments.
Polypedilum (Polypedilum) altkola Kieffer, 1913
Polypedilum alticola Kieffer, 1913. Res. Scient. Voy. Ch. Allaud et R. Jeannel en Afrique orientale (1911-1912) (Dipt.) 1:
22, Type-locality; Kenya, nr Fort Hall [Type(s) not examined].
Polypedilum alticola. Freeman 1958; Bull. Br. Mus. Nat. Hist. Ent. 6: 272.
Material: I d', Saudi Arabia: Asir, Abha, 18° 14'N 42°56'E, 2260 m o.d" 15.VII.1981, W.BUttiker. W (1 slide, alco­
hol), Saudi Arabia: Asir, Adama, 19° 19'N 42°04'E, 1950 m o.d., 23. IX. 1978, W. BUttiker. 1 d' (alcohol), Saudi Arabia: Asir,
Ash Sharayi W., 18°04'N 43° 38'E, 2000 m o.d., 24.-25. IX. 1978, W. Biittiker. 2 d'd' (d'd' and 99 in alcohol), Saudi Arabia:
Asir, Al Dalhan, 18 °oI'N 43 0 24'E, 2180 m o.d., 19.-20. IX. 1980, W. BUttiker. - d'd' (in alcohol), Saudi Arabia: Makkah, Harithi,
21 0 18'N 40018'E, 1910 m o.d., 1984, W.BUttiker. d'd' (in alcohol), same data, 23.-24.V.1985. 99 (in alcohol), Saudi

I
FAUNA OF SAUDI ARABIA 10,1989
I;igs 40-45: 40, Microtendipes umbrosus, wing, 41, 42, Paratendipes nubilipennis, wing, hypopygium, 43, Polypedifum (Pentapedifum)
wittei, hypopygium. 44, Pofypedililm (Pofypedilllm) afticofa, hypopygium. 45, Polypedifum (Polypedifum) nubeculosum, hypo­
pyglUm.
261

264 P. S. CRA:\STON, D. D. JUDD
PoJypediJum (PoJypediJum) nubifer (Skuse, 1889)
Chironomus nubifer Skuse, 1889. - Proc. Linn. Soc. N's.W. (2) 4: 249. Type-locality: Australia, New South Wales, Berowa.
(Lectotype d examined, ANIC).
Polypedilum nubifer. - Freeman 1960; Aust. J. ZooL 9: 707.
Material: 5 dd, Saudi Arabia: Riyadh, Durmah, 24°37'N 46 °07'E, 610 m o.d., 2S.IX.1979, W.Buttiker (2 to NHMB).­
I cJ', Saudi Arabia: Medina, Al Ayda Hafirat, 26 ° 24'N 39° 10'E, 1150 m o.d., 28. IV. 1979, W. Btittiker. - dd (in alcohol), Saudi
Arabia: Eastern Province, Hofuf,. 25 °24'N 49°28'E, 150 m o.d., 7. V.1978, W. Buttiker. - 3 same data, 24.-28. V. 1978. - 1
d, Saudi Arabia: Medina, Wadi Jizl, 26 0 42'N 37° 15'E, 1000 m o.d., 14. XL 1984, W. Buttiker. - several dd (alcohol), Saudi
Arabia: Riyadh, Al Khardj, 24° 15'N 47° 23'E, 410 m o.d., 13. II. 1981, W. Btittiker. - 1 d, Oman, Tahwa nr Sur, 'CDC in goat
pen', 25. XL 1986, R. P. Lane. 1 d, Qatar: Doha, \.-3. IX.1981, W. Buttiker. 2 dd, Qatar, Musimir, 'new lake site', 23. VI.
1981, W. Btittiker.
Hypopygium: fig. 46.
Distribution: Palaearctic region: southern Europe and including North Africa, eastwards
through Egypt, Syria and Iraq to Japan and North Korea. Oriental region: Pakistan, India, Sri Lanka,
Taiwan and Indonesia. Austropacific region: Australia and Micronesia.
Specimens from Turkey (10 km east of Edirne, 3. VlII.1974, P. S. Cranston, BM 1974-482)
appear to be the first record of P. nttbifer from Turkey.
Specimens from Jordan (Azraq Shishan Marsh, 21. Iv'-I1. V.1966, D. S. Fletcher, BM 1966-298)
apppear to be the first record of the species from Jordan.
PoJypediJum (PoJypediJum) tana n. Sp.
Material: Holotype: Ethiopia, Baha, Dar, Lake Tana, 26.1. 1977,JReichholf (ZSM). Paratypes: 2 dd, Saudi
Arabia: Baha, Adnan, 20° 26'N 41 °3 I'E, 1350 m o.d., 21.-23. IX. 1978, W. Btittiker (BMNH).
Description: Polypedilum tana conforms to the generic diagnosis in WIEDERHOLM (1989). Specifically:
Body length 2.5-2.6 mm, wing length 1.4-1.8 mm, antenna] ratio 1.5-1.8, venarum ratio 1.17,
anterior leg ratio 1.5 (n 1).
Anterior tibial apex: fig. 48.
Hypopygium: fig. 47.
Polypedilum lana is differentiated from its relatives in the subgenus Polypedilum by the reduced
fore-tibial scale, while the hypopygium somewhat resembles that of P. convictttm (Walker). The
absence of microtrichia on the superior volsella, and the shape of the volsella at the site of origin of
the medio-Iateral seta are characteristic.
Distribution: Polypedilttm tana is known only from the type-locality in Ethiopia and from
Saudi Arabia.
PoJypediJum (Tripodura) aegyptium Kieffer, 1925.
Polypedilum aegyptium Kieffer, 1925. Bull. Soc. Ent. Egypte 8: 270. Type-locality: (Egypt) Maadi [Type(s) not
examined ].
Polypedilum aegyptium. Freeman 1958; Bull. Br. Mus. Nat. Hist. Ent. 6: 308.
Polypedilum aegyptium. Albu 1980: Fauna Rep. Pop. Rom. 11 (13): 198.
Material: 1 cJ', Saudi Arabia: Makkah, \'Qadi Turabah, 20° 29'N 41 0 15'E, 21. IV. 1980, W. Buttiker. lOman, Tahwa nr
Sur, 'CDC in goat pen', 25. XI. 1986, R. P. Lane.
Hypopygium: fig. 49.
Anterior tibial apex: fig. 50.
Distribution: Palearctic region: Sweden, France, Spain, Romania, North Africa (Morocco to
Egypt); Afrotropical region: Sudan, Niger; Oriental region: India, Burdwan & Pasighat.
Co mm en ts: ALBU (1980) synonymised the Afrotropical species Polypedilum pruina Freeman with
Polypedilum aegyptium. Examination of the type of Polypedilum pruina (BMNH) suggests that P.
pruina may be distinct from P. aegyptium, based on wing venational and anal point structure. How­

L
FAUNA OF SAUDI ARABIA 10, 1989
ever, ALBu (1980: figs 134-136) illustrates these extremes of variation from Romanian material that
she assigned to P. aegyptium. Saudi Arabian material falls within this variation.
As with much material from this Arabian collection, the wing pattern is very weak, but appears to
resemble ALBu's (1980) fig. 134a rather than the more typical'P. pruina' of her fig. 134b.
Stictochironomus puripennis (Kieffer, 1921)
Polypedilum puripenne Kieffer, 1921b. - Ann. Soc. Scient. Brux. 41: 97. Type-locality: (South Africa) Transvaal (Type not
examined).
Stictochironomus puripennis. - Freeman 1958; Bull. Br. Mus. Nat. Hist. Ent. 6: 308.
Material: 2 Saudi Arabia: Medina, AI Ayda Hafirat, 26"24'N 39°1O'E, 1150 m o.d., 28.IV.1979, W.Blittiker (I to
NMHB, I to ZSM). I d Saudi Arabia: Medina, Khaybar, 25°36'N 39° 16'E, 680 m o.d., 26.IV.1979, W.Blittiker. I d,
Saudi Arabia: Makkah, Wadi Minsah, 20 0 31'N 40 0 40'E, 550 m o.d., 7.-8.IV.1983, W.Blittiker. 1 Saudi Arabia: Baha,
Jebel Ibrahim, 20° 2S'N 41 0 IrE, 1540 m o.d., 10.-11. IX. 1983, W. BUttiker. 2 dd, Saudi Arabia: Baha, Wadi Ilyab, 20 0 07'N
40 0
57'E, 160 m o.d" 3.11.1984, W.Blittiker. - 1 d, Oman: Nizwa, 14.X1.1986, R.P.Lane; 1 d, Oman: Tahwa nr Sur, 'light
trap in goat pen', 25.XI.1986, R. P. Lane.
Scutal tubercle: fig. 51.
Hypopygium: fig. 52.
Distribution: widespread in Afrotropical region from Sudan southwards. These records constitute
the first outside the Afrotropical region.
Genus unknown
Material: I Saudi Arabia: Makkah, Wadi Maraum, 22" 16'N 39° 44'E, 280 m o.d., 3.-4. V. 1984, W. Btittiker. 2 dd,
Saudi Arabia: Gizan, Wadi Damad, 17° 17'N 43°06'£, 800 m D.d., 24.IX.1981, W.Btittiker.
Hypopygium: fig. 53.
Distribution: This species is known only from two localities in relatively low altitudes in southwestern
Saudi Arabia.
Comments: This species, represented by three specimens from two localitites in Saudi Arabia,
has a highly distinctive male hypopygium, unlike any other described Chironominae. It will not satisfactorily
key to genus, and without the immature stages, it would be premature to speculate on its
relationships. For this reason we refrain from naming the taxon, but illustrate the hypopygium and
offer the following diagnostic comments:
Head: Antenna with 11 flagellomeres, tapering apically to blunt point; ratio of c. 2.0. Frontal
tubercles minute. Two pairs of ocelli in relatively large ocellar triangle. Vertical setae biserial. Palp 5
segmented, with two fine sensilla chaetica subapically on segment 3, without a pit.
Thorax: with narrow antepronotallobes, probably narrowly separated medially anterior to apex
of non-extended scutum. Acrostichals long, more or less biserial, starting at anterior of scutum; dorsocentrals
fewer, uniserial; 3-4 prealars, about 10 uniserial scutellars; scutal tubercle present.
Legs: Anterior tibial scale bluntly pointed, mid and hind tibial combs proximated, each bearing a
short spur; tarsomeres 4 and 5 subequal, pulvilli indistinct, about half length of claw. Sensilla chaetica
uniserial in apical half of mid tarsomere 1 and apical two-thirds of hind tarsomere 1.
Wing: with moderate to fine punctation, with R2+3 running midway but ending at one third distance
between R] and R4+5' without any costal extension. Squama variably damaged, but no seta visible
and is likely to be bare.
Abdominal setation: moderately dense but not organised into any discernible pattern either
on tergites or sternites. There is a slight indication of darker tergal bases.
Hypopygium (fig. 53): The setae dorsally on the anal point are quite characteristic and may be
unique. The superior volsella shape, with microtrichia and setae only on the median half, somewhat
resembles some Dicrotendipes, but there is no obvious curvature of the inferior volsella. There is no
indication of any median volsella.
265

lit:
FAUNA OF SAUDI ARABIA 10,1989
Figs 58-62: 58, Tanytarsus IlOmi, hypopygium. 59, Tanytarstis mcmillani, hvpopygium. 60, Tanytarsm paliiduLus, hypopygium. 61,
62: TanytarsU5 trijidm, 61, 62, median volsella (appendage 2a).
62
269

FAUNA OF SAUDI ARABIA 10, 1989
Hypopygium: fig. 6l.
Median volsella (appendage 2a): fig. 62.
Distribution: T trijidl1swas known previously only from the Afrotropical region, from Nigeria
(Kankiya and Zaria) and Burkino faso (Upper Volta) (fREEMAN 1958). These specimens comprise the
first records from outside the Afrotropical region.
Comme n ts: This species has a very distinctive median volsella (appendage 2a), resembling that
of the western Palaearctic T excavat115 group (REISS & FITTKAU 1971). However, T trifid115 differs
from all the included species in the T excavatl1s group by possessing combs and spines on the anal
point.
Virgatanytarsus arduennensis (Goetghebuer, 1922)
Tanytarsus arduennensis Goetghebuer. 1922. - Ann. bioI. lacustre 6: 43. Type-locality: Belgium.
VirgatanytarsttS arduennensis. - Pinder 1982; Spixiana 5: 32
Tanytarsus subrcjlexens Freeman, 1955a. - Expl. Pare Nat. Albert Miss. Witte 83: 37. Zaire, Vitshulllbi
(Holotype d' examined, MRAC). n. syn.
Tanytarsus subrejlexens. - l'reeman, 1958: Bull. Br. Mus. Nat. Hist. Ent. 6: 344.
Material: 2 Saudi Arabia: Makkah, Khoda, 20° 22'N 41 ° 18'E, 1890 m o.d., 19. IX. 1982, W. BUttiker. 2 Saudi
Arabia: Eastern Province, Hofuf, 25°24'N 49°28'E, 150 m o.d., 28. V.1978, W. BUttiker. lOman, Tahwa nr Sur, 'CDC in
goat pen', 25.XI.1986, R.P.Lane.
Hypopygium: fig. 63.
Distribution: Palaearctic region: widespread from Britain to Finland, eastwards through middle
Europe to Yugoslavia, Turkey and Palestine. l'\ew records from Turkey are: 1 d, Sivas, Gurun,
15.VIII.1974; 1 d, Konya, 28 miles SWKonya, 3r42'N 32°24'E, 13. VIII. 1974, both p.s.Cranston,
BM 1974-482. Afrotropical region: Ethiopia, Ghana, Nigeria and Zaire.
Comments: Species delimitation in Virgatanytars115 is extremely difficult. KUGLER & REISS
(1973), in revising the Tanytarsus triangularis group, which became the genus Virgatanytarsl1S (PINDER
1982), recognised seven species from the western Palaearctic and Afrotropical regions. Since then,
examination of more extensive material from a wider geographical area, including the present study
of material from Arabia and environs, gives indications that species variability is higher than previ-
Figs 63, 64: 63, Virgatanytarms arduennensis, hypopygium. 64, Virga ta nytarsus nigricornis, hypopygium.
271

272 P. CRANSTON, D, D. JUDD
ously understood and some synonymy is likely. The most radical suggestion would be to recognise
only two species, namely v. albisutus Santos-Abreu (syns T arduennensis Goetghebuer, T richmondensis
Edwards, T maroccanus Kugler & Reiss, T subre./lexens Freeman & T triangularis Goetghehuer)
possessing elongate rods on the anal point, and v. nigricomis Goetghebuer (syn. T hu/ensis Kugler &
Reiss), with short lobe-like rods on the anal point. However, although we do propose some new synonymy
(see below) we refrain from formally proposing the above solution because we have not
examined the available immature stages, particularly the pupae, nor have we seen material that lies
within the variation of V. triangularis.
FREEMAN (1958) in re-describing T subre./lexens, noted the similarity to the Palaearctic T re./lexens
Edwards [a junior synonym of V. triangularis (Goetghebuer)] and T richmondensis Edwards [a junior
synonym of V. arduennensis (Goetghebuer)]. Examination of additional material shows that the slight
differences observed by FREEMAN (1958) no longer permit recognition of two species. The characters
suggested by KUGLER & REISS (1973) for discriminating the two species was based upon examination
of a single specimen of T subre./lexens more extensive Afrotropical material shows that there is
greater variation than they were aware of. The relative length of the digitus (appendage la) to the
superior volsella (appendage 1) is particularly variable, but in no specimens examined does the digitus
reach the median margin of the superior volsella.
Therefore T subre./lexens falls as a junior synonym of v. arduennensis (Goetghebuer).
Virgatanytarsus nigricornis (Goetghebuer, 1935)
Tanytanus nigricornis Goetghebuer, 1935. Rev. Zool. Bot. Afr. 26: 398. Type-locality: (Zaire) Belgian Congo, Kamande
(Holotype ef' examined, MRAC),
Tanytanus hulensis Kugler & Reiss, 1973. Ent. Tidskr. 94. Type-locality: Palestine, Lake Hula (tlolotype para-
types ef'ef' examined, ZSM). Syn. nov .
.Material: 2 dcJ, Saudi Arabia: Baha, Wadi Diyhan, 19° 48'N 41 0 36'E, 1350 m o.d., 7. III. 1984, W. BOttiker (NHMB).
Ief', Saudi Arabia: Asir, .'\lamas, 19° I (N 42 0
19'E, 2380 m o.d., 11.IV. 1980, W. BUttiker.
Hypopygium: fig. 64.
Distribution: Palaearctic region: Palestine. Afrotropical region: widespread on the east of the
continent, from Yemen southwards to Transvaal. DEJoux (1984) provides a first west African record
from Guinea.
Comments: T hulensiswas established by KuGLER & REISS (1973) for a species showing a close
relationship to the Afrotropical T: nigricomis Goetghebuer. Six differentiating characters were listed
for T hulensis (T nigricomis in parentheses): 1) thorax with dark brown vittae (light green); 2) wing
cells cu and an with many macrotrichia (few); 3) tibial combs with spur (without); 4) maximum of 3
sensilJa chaetica on tarsomere 1 of mid leg (15-18 sensilla chaetica); 5) superior volsella (appendage
I) without microtrichia dorsobasally (with a distinct patch); 6) median volsella (appendage 2a) shorter,
apically broader (longer, not so apically distended). KUGLER & REISS (1973) suggested that speci­
mens of T nigricornis from south of the Sahara might well be found to belong to T hulensis but did
not examine them. Slide preparations of all BMNH Afrotropical 'T: nigricornis' shows that all would
belong to an expanded definition of T huie1l5is, with greater numbers of sensilla chaetica (up to nine
in Yemen specimens) and a much more variably shaped and lengthed median volsella. Body colour
also varied but the intensity of thoracic vittae did not correlate with genitalic features. No Afrotropi­
cal specimens possessed microtrichia on the superior volsella; the only individual in BMNH collections
clearly identical in all features with' T nigricornii in the sense of KUGLER & REISS (1973) was
from Palestine. Further specimens from this country preserved in ZSM, determined as T nigricornis,
have been examined. It is noticeable that specimens that are closest to the concept of T hulensis are
early emerging (said by KeGLER & REISS [1973] to be univoltine), and must have developed in the

fAUNA or SAUDI ARABIA Ie, In9
generally cooler waters than available to 'T. nigricornis'. It is conceivable that the observed differences,
particularly those involving pigmentation and possible allometry, are related to seasonal differences.
The immature stages associated with each species are shown by KUGLER & REISS (1973) to be
very slight.
Saudi Arabian specimens are intermediate between the two KUGLER & REISS (1973) defined species,
being pale, with few microtrichia on the wing cells cu and an, with small but distinct spurs on the
combs, with long median volsella, and, crucially, 12-14 sensilla chaetica on tarsomere 1 of the midleg.
Examination of Goetghebuer's type of T. nigricornis confirms it to be identical TO KUGLER &
REISS'S (1973) T. hulensis and the senior name, T. nigricornis is applied here. It remains a possibility
that specimens possessing microtrichia on the superior volsella, identified incorrectly as T. nigricornis
by Kugler & Reiss (1973), may warrant specific status. Since no specimens possessing this feature
have been found in Saudi Arabia or in our re-examination of Afrotropical and Yemeni BMNH specimens,
we refrain from commenting further on this putative taxon.
DISCUSSION
Ecology
The immature stages of Chironomidae can dominate freshwater ecosystems in terms of both
absolute numbers (biomass) and species diversity. The responsiveness of the chironomid community
to environmental disturbance, such as pollution, has led to recognition of their usefulness as indicators
of such stress. Associated with these studies is the recognition of the need to be able to accurately
identify the immature stages. In some parts of the northern Palaearctic region, and to an extent in
North America, it is possible to make species level identification. Outside these restricted areas, identification
of the immature stages is difficult, if not impossible. Saudi Arabia is such a case. A first
requirement is to gain some understanding of the fauna based upon adult midges. The present study
is the first to endeavour to list the Chironomidae occurring in the region, and is based virtually exclusively
upon adult males - information on the immature stages is minimal.
Adult Chironomidae collected at light are probably rather poor indicators of the ecology of the
associated larvae. The ability to fly some distance to light from the natal site, and the complexity of
microhabitats within that radius, means that we will refrain from all but the most general speculations
concerning ecology.
What we know of larval biology of the African species that dominate the Arabian fauna is that
the vast majority are eurythermic, able to tolerate high water temperatures. This is true of species
from lotic (running) and lentic (standing) waters, both of which appear to be well represented
amongst the collecting sites in Saudi Arabia. The conventional reason why the subfamilies Chironominae
and Tanypodinae dominate in these latitudes is their eurythermy, relative to the generally more
cold adapted subfamilies such as the Podonominae and Orthocladiinae. CRANSTON et a1. (1987) challenge
the orthodoxy of the cold stenothermic origins of the Chironomidae, but it is clear that subsequent
radiation at subfamilial level is quite strongly associated with differential abilities to thrive in
various thermal regimes. The determining factor is unlikely to be temperature alone but the availability
of dissolved oxygen, which is much reduced in the high water temperatures encountered in low
latitudes, particularly in standing waters at low elevations. The widespread occurrence of haemoglobin
in larval Tanypodinae and Chironominae must assist in gaining an advantage over the Ortho­
273

274 P. S. CRANSTON, D. D. JUDD
cladiinae lacking haemoglobin, in oxygen poor waters (CRANSTON 1988). Interestingly, the Orthocladiinae
found in Arabia are predominantly of northern origin, or, if present in the Afrotropical region
(Paratrichoc!adius micans), predominantly with montane distribution. All Arabian Orthocladiinae are
found at altitudes in excess of 600 metres in the north of the country and at some of the highest altitudes
of up to 2,500 m o.d. in the peninsula when present in the south. Although it is difficult to separate
the effects of historical events from current ecological conditions, in this case it seems likely that
the Orthocladiinae are restricted to cooler water temperatures than are many of the T anypodinae and
Chironominae.
Reports of some Chironomidae causing nuisance by their sheer abundance in polluted waters are
increasing throughout the world (CR.I\NSTON 1988) and Arabia is no exception. Kiei/erufus disparifis
was reported to be creating a nuisance when flying around Jeddah Sewage Works as was Chironomu5
cafipterus at Zakhar Alain, U.A.E. (G.B. WHITE pers. comm.). Although adult Chironomidae are
known to be potent environmental allergens, notably in the Sudan (CRANSTON et. al. 1981) there are as
yet no reports of such medical symptoms in Arabia.
Biogeography
Although Arabian adult Chironomidae collections do not allow us to assess the regional value of
this family in indicating aquatic ecological conditions, they are valuable in faunistic studies and
appear to be eminently suitable as a tool for analysis of biogeographical relationships. Such an exercise
depends upon an adequate knowledge of the regional fauna and firstly it must be asked how well
we know the fauna of Saudi Arabia and adjacent countries?
The Arabian Chironomidae fauna revealed by collections from 74 localities in Saudi Arabia, 2 in
Oman,2 in Qatar, 1 in UAE and 1 in Dubai, comprises 53 species in 3 (of 8 worldwide) subfamilies.
Appropriate comparisons are with the fauna of Syria, with 71 species (REISS [1986] supplemented by
this study) and Ethiopia with 56 species (FREEMAN & CR.I\NSTON [1980]; HARRISON [1987]). The subfamily
composition in Ethiopia is identical to that of Arabia, whereas in Syria, with a predominantly
Palaearctic fauna, the additional Holarctic subfamily Prodiamesinae occurs. Turkey, with a similar
but even more clearly Palaearctic fauna, has 103 species (REISS [1985], supplemented by this study)
and both Diamesinae and Prodiamesinae are present. The Lebanon, where the rivers Orontes and
Litani were intensively studied by MOUBAYED & LAVILLE (1983), has a fauna of at least 140 species, of
which the greatest proportion are Palaearctic in distribution. In contrast, Palestine, particularly the
lakes Tiberias and Hula, have a more restricted fauna of about 50 recorded species (notably documented
by KUGLER [1966], KUGLER & WOOL [1968]) which, although originally described as predominantly
Afrotropical, must now be seen as basically Mediterranean, or even eremic (see below). In contrast
to these relatively well studied areas, no Chironomidae have been reported from Somalia, and
few from Iraq and Iran. In terms of understanding endemicity, the lack of information from the People/
s Democratic Republic of South Yemen (Aden) is regrettable, but may be compensated for by
examination of historic collections from North Yemen and contiguous areas of Saudi Arabia. Despite
these reservations, we believe that the Arabian fauna revealed in CR.i\NSTON (1989) and this study is
adequately comparable with other regional studies and that collecting artefacts are unlikely to affect
our biogeographic assessment.
Since our comparisons are largely with the Afrotropical fauna, it is necessary to establish the
depth of this knowledge. Some countries were ratherly poorly collected when FREEMAN & CRANSTON
(1980) catalogued the regional fauna. However, despite additional study, there have been relatively

FAUNA OF SAUDI ARABIA 10, 1999
has a more restricted distribution in the same latitudes (fig. 65). Thus, in the Chironomidae, we find
four endemics reflecting LARSEN'S (1984) dominant SW Arabian Rhopaloceran endemism and two
endemic species from his restricted Oman and desert zone.
LARSE!': (1984) regretted the limitation of his ability to make comparisons of Rhopalocera endemicity
with that of other arthropod groups, but, from fragmentary data, suggested that butterfly
endemism was at a lower taxonomic level than in other comparable groups. In reviewing previous surveys,
LARSEN (1984) found only Odonata to have no endemic Arabian species, compared with high
endemism in Bombyliidae (Diptera), Tenebrionidae (Coleoptera), Orthoptera and Lepidoptera: l\1acroheteroptera.
However, more recent study of the Odonata has shown endemism, at least in the
genus Pseudagrion (SCHNEIDER 1987). The disparity between the findings of other scientists and his
own prompted LARSEN (1984) to an extensive discussion of the reasons for the lower endemism in
Rhopalocera.
Our findings of a remarkably similar low level of endemism to that of the Arabian Rhopalocera
provoke some comments on LARSE!':'S (1984) discussion. LARSEN suggested three a priori reasons why
one might expect a low level of endemicity: 1) high vagility, reducing isolation, 2) visual courtship acting
to stabilize taxonomically important colour patterns, and 3) diurnal behaviour leading to intolerance
of environmental fluctuation, and hence leading to extinction. All three explanations were felt to
apply to other groups with apparently restricted endemism. Furthermore, in an earlier section, artefacts
of collecting were raised (but dismissed) as a general criticism of the reality of the patterns of
endemicity detected.
How do these relate to the Chironomidae? Firstly, there is little doubt of the vagility of Chironomidae:
they rank second only to aphids in the aerial plankton, yet they are not known to undertake
the characteristic directed migration of some butterflies. lIowever the vagility of organisms is overrated
in discussions of the role of dispersal in the composition of extant faunas - it is the ad hoc
explanation of any incongruence in distribution, and is intimately tied up with untenable ideas of
"centres of origin". That dispersal has occurred is beyond doubt COOPE (1979) and GENTRY & SUT­
CLIFFE (1981) amongst others have conclusively demonstrated alteration of ranges amongst Quaternary
animals in relation to Pleistocene climatic changes, including expansions and contractions of glaciations.
Subsequent to cladistic analysis, WILLASSEN & CRl\.NSTON (1986) concluded that the present
distribution of Diamesa (Chironomidae) in montane East Africa must have resulted from dispersal
from Palaearctic ancestors. However, such dispersalist scenarios depend on newly created vacant habitats
- in these examples, the postglacial terrain of the Holarctic region and the glaciated mountains
of East Africa respectively. The dispersalist argument is that organisms from one region diffuse/ disperse
into another, where related organisms already occur in presumed harmony with their environment,
and establish themselves there. Dispersal, the arrival of an organism into a new area, is undisputed
establishment by displacement, or incorporation into a new ecosystem other than those
anthropogenically disturbed, is the important factor. Vagility, as a taxon characteristic, does not seem,
in principle, to be correlated with a ready ability to establish and retain a viable population in preexisting
ecosystems, the criterion required if dispersivity is seen as a major factor in the historic structuring
of communities. A test that vagility is little related to distribution patterns (other than those
strongly modified by anthropogenic disturbance) must be the observation that highly vagile organisms
such as Aves, Chironomidae and Rhopalocera are not cosmopolitan but show distributions that
are highly correlated with earth history (e.g. CRACRAFT [1982J - Australian birds; BRUNDIN [1966J
Australian Chironomidae; KITCHING et. al. [1987] - Rhopalocera, Danainae, Idea; VANE-WRIGHT
[pers. comm.J - Rhopalocera of Sulawesi). CROIZAT (1958) made the point more than quarter of a
century ago when he contrasted the gross differences in dispersivity between hawks and land snails
and the equivalence of their actual restricted distributions.
277

278 P. S. CRANSTO", D. D. JUDD
LARSEN'S (1984) second point, that in Rhopalocera, highly developed courtship leads to stabilizing
selection on a primary taxonomic character, colour pattern, does not seem to apply to the Chironomidae.
The specificity of male chironomid swarm site, in conjunction with highly species-specific male
genitalia (that correlate well with Immature stage taxonomy) suggests that species discrimination in
the Chironomidae differs from that of Odonata and Rhopalocera; yet the same pattern of endemism
is discovered.
LARSEN'S (1984) third point, that endemism is higher in those groups that react to climatic deterioration
by "adapting", rather than going extinct, is essentially untestable. Sympathising with BDITON'S
(1987) sceptical views on adaptationist hypotheses, we are unable to comment on this explanation.
The possibility of collection artefacts resulting in a biased assessment of endemism has already
been discussed above. Like LARSEN (1984), we believe that the collections made in Arabia accurately
reflect the studied fauna in comparison with those of other parts of the Near and Middle East. No
doubt, the trapping method utilised exerts a bias and more intensive collecting by a specialist would
increase the list, but the same would apply to the faunas of Ethiopia and Syria. It is pertinent here that
an increase in knowledge of the barely known faunas of Somalia and Iraq would be likely to diminish
the already low level of Arabian endemism.
In summary, endemism in Saudi Arabian Chironomidae is low, and of the same order as that
found for Rhopalocera by LARSEN (1984). Our view is that this is not an artefact but a genuine observation
that requires explanation in historical terms.
Widespread taxa
Since historical biogeography demands a knowledge of sister group relationships, does this imply
that without a hypothesis of phylogeny, biogeographic researches are fatally flawed? Are all faunistic
studies such as this simply descriptive, able to contribute only in a narrative manner to theories of faunal
relationships? While accepting that the rig our needed to put a time frame on branching events can
only be obtained through cladistic analysis, followed by production of area cladograms, congruent
patterns still can be detected and related to earth history events in the absence of a hypothesised
phylogeny.
The classic case of faunal patterns being self-evident without a theory of cladistic relationship is
the detection of the world's zoogeographic regions. This prevalence of faunal similarities and disjunctions
was well understood by SClATER (1858) and WALLACE (1876) and led to the delimitations of faunal
regions (for example, by SCLATER & SCLATER [1899]) that were readily accepted, with disagreement
only concerning exact location of the boundaries. This clear demarcation of biogeographic
regions is strong evidence for congruence of repeated distribution patterns and is most parsimoniously
interpreted as being produced by biotic responses to identical vicariance events. Of course, such
ideas flowered fully in the extensive studies of CROIZAT (e.g. 1958) and summarised, with the additional
non-Croizatean demand for monophyly of the treated groups, by CROIZAT et al. (1974).
In the Chironomidae, as with the Rhopalocera studied by LARSEN (1984), the dominant distribution
patterns are so evident as to be almost mundane. The predominant faunal elements are Afrotropical
in their non-Arabian distribution (tab. 2). Thus, excluding the six endemic species, 49% of Arabian
Chironomidae species are otherwise found exclusively in the Afrotropical region. When Afrotropical
species that also occur contiguously in the Palaearctic region and/or Oriental regions are
included, then this proportion rises to 75% of the fauna. This predominance and proportion of Afrotropical
species is very similar to that found by LARSH; (1984) for the Arabian butterflies (Rhopalocera).

LAcUNA OF SAUDI ARABIA 10,1989
Table 2: Arabian Chironomidae otherwise restricted to Afrotropical region.
Procladius apicalis
Procladius brevipetiolatus
Procladiu5 reidi
Conchapelopia tri/ascia
Lania rutshuruiell5is
Lania teesdalei
Paramerina sp. 1 sensu LEHMANN
Cricotopus albitibia
Chironomu5 seychelleanus
Cryptochironomus diceras
Dicrotendipes freemani: N.E. + Seychelles
Dicrotendipes sudaniws: Sahelian
Kie./Jerulus disparilis: Sahelian
iHicrotendipes umbrosus
Paratendipes nubilipennis
Polypedilum wittei
Po(ypedilum lana: N.E.
Stictochironomus puripennis
Cladotanytarms pseudomanCIIs
Cladotanytarsus reductHs
Tanytarsu5 mcmillani: Sahelian
Tanytarslis pallidulus
Tanytarsw tri/idus
Amongst the Afrotropical Tanypodinae, only P. apicalis is widespread; the other species are
restricted to the west, predominantly the southwest and at middle to high altitudes (fig. 66). The sole
Afrotropical Orthocladiinae, C. albitibia, has similar distribution, being found only at 17° S. Within
the Afrotropical Chironominae, all species occur in the west to southwest. Of the 15 species, 9 are
restricted to this area, 4 extend eastwards to Hofuf and Oman, and 2 widespread species have northern
extensions into Jaw£. Thus the dominant distribution pattern within Arabia of otherwise Afrotropical
species (fig. 66) is the occurrence at altitudes from 100 m o.d. to over 2000 m in western to
..' MFOINA
RIYADH
IRAN
U.A.E.
PROV,NCE
55"E
\
/
/
('
j,
J' \
i I
o Larsia rutshuruiensis
o Cricotopus albitibia
f'. Paratendipes nubilipennis
• Stictochironomus puripennis
• Cladotanytarsus reductus
Fig. 66: Distribution in Arabia of representative Chironomidae otherwise restricted to Afrotropical region; Larsia nltshuruiensis,
Cricotopus albitibia, Paratendipes nubilipennis, Stictochironomus puripennis, Cladotanytar5Hs reductus.
•
279

FAUNA OF SAUDI ARABIA 10, 19B9
nothing more than "noise" induced by the ecological equivalent of the systematists' homoplasy - i. e.
the result of random diffusion from a region of origin into a proximate area. LARSEN (1984) had similar
doubts, believing that the category may be a "catch-all" for taxa that do not fit traditional zoogeographic
categories. We do not believe that our discovery of such a similar grouping denies or sustains
the existence of an Eremic category of zoogeographic subregion and agree with LARSEN's (1984:
133) reservations on the matter. However, suggestions that these species may reflect a purely ecological
response to eremic (e.g. xeric) conditions might be tested by the discovery of such a grouping in
the Chironomidae. There could hardly be a greater disparity between ecologies of the Rhopalocera,
reliant (often very specifically) on terrestrial plants for larval development, and the Chironomidae
with aquatic immature stages. The existence of Chironomidae that shared mechanisms for resisting
the erratic drying out of eremic water bodies might be considered to be an ecological congruence with
the Rhopaloceran limitation by xeric vegetation, but we have no evidence of such chironomid adaptations
in the region. Thus the existence of at least some apparently eremic Chironomidae indicates that
this zone is unlikely to the ecological phenomenon proposed, but may indeed reflect a genuine historical
pattern.
In five distribution categories discussed above, there is a quite remarkable congruence of the proportions
of species allocated to each, by LARSEN (1984) for Rhopalocera and ourselves for the Chironomidae.
Disparities in ecologies between these groups is a strong indication that this congruence is
not prima facie due to similarity of responses to modern day environment and climatic factors. These
repeated overall higher category patterns (themselves composed of repeated species level patterns) are
real, despite the paucity of cladistic information, and require explanation in terms of earth history
driving common distributions in both Rhopalocera and Chironomidae.
Earth History
Although BALL'S (1975) warning to biogeographers to avoid rationalising their data to conform to
current geological opinion is well taken, the congruence between regional earth history and faunal
distribution pattern is evident.
Taking our data from the excellent palaeomaps of OWEN (1981), HOWARTH (1981) and ADAMS
(1981), the relative position of Arabia can be traced from the Triassic, 240 my bp to the present. Palaeontological
and cladistic biogeographic analysis indicates that the Chironomidae were present and
diversified in the Upper Jurassic (c. 150 my bp) (CRANSTON et al. 1987) and a starting point in the Triassic
is thus appropriate.
At this time the Arabian peninsula constituted the northeast corner of the African plate. The African
plate was very close to western India and what was to become the northern and northeastern part
of Arabia was beneath the Tethys Sea. Throughout the Lower Jurassic there was little change in the
proportion of Arabia that was above the ocean, but by the Kimmeridgian Upper Jurassic (145 my bp)
a rise in sea level left only a southwestern third of the country above sea level, as the southern part
became inundated by an expanding Tethys. The Indian plate remained proximate, but separated from
the NE corner of the African plate by the epicontinental sea.
In the ensuing 80 my, the African plate continued to move northwards and its position relative to
India remained more or less constant. The above ocean part of Arabia remained restricted to a southwestern
section, with the north and east immersed. It was only at the start of the Tertiary (65 my bp)
that Africa rotated somewhat to give Arabia a more North-South axis and increased proximity to
Eurasia. India moved away northeastwards. Low-lying Arabia (northern and eastern parts) remained
beneath the contracting Tethys, particularly during the maximum extent of seas in the Palaeocene. At
283

284 P. S. CRANSTON, D. D. JUDD
low sea during the late Oligocene, ADAMS (1981) suggests the Tethys may have been interrupted by
connection from Turkey to the Arabian part of the African plate,
In the Miocene (c. 25 my bp) the anticlockwise rotation of Africa had pushed Arabia to virtual
connection with Asia, with the Tethys reduced to a remnant connecting the proto-Mediterranean to
the Indian Ocean. At this time virtually all of modern Arabia was above sea level. Meanwhile, after
over 200 my of connection between Arabia and the rest of the African plate, rifting that commenced
in the late Oligocene (24-20 my bp) and continued through the Miocene had led to the formation of
an incomplete Red Sea. ADAMS (1981) cites sources suggesting a complex history for the Red Sea,
with a Miocene connection only to the proto-Mediterranean, replaced in the Pliocene by an intrusion
of the Indian Ocean, with the northern connection to the Mediterranean closed by uplift.
ADAMS (1981) also discusses briefly the probability of islands occurring in the Miocene Tethys,
which might have afforded avenues for faunal exchange between the African plate and the composite
and often fragmentary plates to the north and east.
Earth history and present biota
How does this geological history relate to the present day faunal distribution? firstly, the Arabian
distribution of the Afrotropical species listed in tab. 2, coincides with the area that has been an
integral part of the African plate and was never inundated during the maximum Tethys sea level. The
precise delimitation of this area is unclear. fluctuations of sea level especially in the Tertiary, outward
diffusion of Afrotropical species and variable incursion of non-African elements naturally will have
blurred any strict demarcation of the limits to the Afrotropical faunistic dominance. However, it is
evident that the montane areas of western and southwestern Arabia support an undeniably Afrotropical
fauna. The recency of the Red Sea rifting, and the ineffectiveness of the resultant barrier, is indicated
by the one putative case of vicariant sister species discussed above.
Given an Afrotropical dominance, it is pertinent to examine whether there are any Gondwanan
affinities that might be used to date the age of some faunal elements. The modern African fauna has
relatively few recognised Gondwanan elements and these are restricted, so far, to the subfamilies Podonominae,
Aphroteninae and Diamesinae, occurring in the montane southern part of the continent.
Other probable Gondwanan elements, as yet little studied, include genera of the Orthocladiinae and
particularly the Chironominae, notably Conochironomus and Skusella. None of these occur in Arabia,
nor in the northern two-thirds of the continent.
The areas of current-day Arabia that support a Palaearctic dominated fauna were beneath the
Tethys, at least until the final lowering of sea levels in the late Palaeogene. Palaearctic species must be
seen as having dispersed, or, probably more correctly, to have diffused, from newly contiguous area
of the Palaearctic. The present-day distribution of these species is rather haphazard. Many are present
in the low-lying areas of northern Arabia, extending eastwards, while others are montane, presumably
tracking ecologically suitable conditions. Two species occur in lowland coastal southwestern
Arabia, a distribution that doesn't easily admit explanation.
Examining the distribution of Palaeotropical species, the dominant distribution is in Arabia that
is known to have been inundated by the Tethys. Once again, it is likely that these are dispersive,
weed like, species that have the ability to colonise new habitats created as northern Arabia rose from
the Tethys, in this case, either from the now proximate Asia or from the Palaearctic. Tests to detect
the origin of these widespread taxa can be made only from cladistic hypotheses of relationships.
However, it is clear that these taxa must have dispersed at a relatively recent date from newly proximate
sources in the western Palaearctic and Orient. The role of India is quite uncertain - as with

FAUNA OF SAUDI ARABIA 10, 1989
Archaeochlus (CRANSTON et al. 1987) the lack of reliable information from Gondwanan southern and
western India is a major deterrent in cladistic biogeographical studies,
OWEN (1981) questions the constant dimension globe used by most geologists and suggests an
expanding earth model is more appropriate, In the region studied here, the difference between the
two is the nature and extent of the Tethys, In a constant dimension model, the Tethys is a major oceanic
barrier between the African plate and the Palaearctic. In OWEN'S (1981) espoused expanding earth
model, this wide Tethys is seen .as an artefact of continental projection onto a constant dimension
globe. On a historically smaller globe, the Tethys is seen to be, at most, a shallow epicontinental sea.
Faunistic studies in Arabia may help eventually to provide some evidence concerning the role of
the Tethys in the development of the Arabian fauna, particularly in assessing its efficacy as a barrier
to dispersal into Arabia from the northern continental mass. However, here it is clearly necessary to
have a cladistic hypothesis of taxic relationships on each side of the Tethys. It is possible that the two
lowland endemic Arabian Chironomidae reflect isolates on Tethyean islands offshore from the African
plate, now united in Arabia. Again, these are no more than speculations in the absence of a cladistic
hypothesis.
SUMMARY
The chironomid fauna of the Arabian peninsula is shown to consist of 53 species, comprising 10
members of the subfamily Tanypodinae, 5 Orthocladiinae and 38 Chironominae, of which 13 are
Tanytarsini.
The following nomenclatural changes are made:
Bryophaenocladius paraproductus (Orthocladiinae) from Saudi j\rabia and Polypedilum (Polypedilum)
tana (Chironominae) from Ethiopia and Saudi Arabia, are described as new to science.
One new combination is erected, Kie./ferulus disparilis (Goetghebuer, 1936), based upon the first
description of the immature stages. Two new synonymies are proposed: Tanytarsus subrejlexens Freeman,
1955 is a junior subjective synonym of Virgatanytarsus arduennensis (Goetghebuer, 1922) and
Tanytarsus hulensis Kugler & Reiss, 1973 a junior subjective synonym of Virgatanytarsus nigricornis
(Goetghebuer, 1935).
Besides the new species discovered in the region, the following species described previously from
elsewhere are recorded for the first time from Saudi Arabia. These are: (subfamily Tanypodinae) Procladius
(Holotanypus) apicalis (Kieffer, 1918), Procladius (Holotanypus) brevipetiolatus (Goetghebuer,
1935), Procladius (Psilotanypus) reidi Freeman, 1955, Ablabesmyia (Ablabesmyia) longistyla Fittkau,
1962, Ablabesmyia (Ablabesmyia) monilis (Linnaeus, 1758), Conchapelopia trifoscia (Freeman, 1954),
Larsia rutshuruiensis (Goetghebuer, 1935), Larsia teesdalei (Freeman, 1955b), Paramerina vaillanti Fittkau,
1962; (subfamily Orthocladiinae) Cricotopus (Cricotopus) albitibia (Walker, 1848), Paraphaenocladius
impensus (Walker, 1856), Paratrichocladius micans (Kieffer, 1918b), Psectrocladius limbatellus
(Holmgren, 1869); (subfamily Chironomidae) Chironomus calipterus Kieffer, 1908, Chironomus seychelleanus
Kieffer, 1911, Cryptochironomus diceras Kieffer, 1923, Cryptochironomus rostratus Kieffer,
1921 a, Dicrotendipes freemani Epler, 1988, Dicrotendipes peringueyanus Kieffer, 1924, Dicrotendipes
sudanicus Freeman, 1957, Kie./feruius disparilis (Goetghebuer, 1936), Microchironomus lendli (Kieffer,
1918b), Microchironomus tener (Kieffer, 1918a), Microtendipes umbrosus Freeman, 1955a, Paratendipes
nubilipennis Freeman, 1957, Paratendipes nudisquama (Edwards, 1929), Polypedilum (Pentapedilum)
wittei Freeman 1955a, Polypedilum (Polypedilum) alticola Kieffer, 1913, Polypediium (Polypedilum)
nubeculosum (Meigen, 1804), Polypedilum (Polypedilum) nubifer (Skuse, 1889), Polypedilum (Tripodura)
aegyptium Kieffer, 1925, Stictochironomus puripennis (Kieffer, 1921 b), Cladotanytarsu5 pseudomancus
285

286 p, S, CRANSTON, 0,0, JUDD
(Goetghebuer, 1934a), Cladotanytarsus reduclus (Freeman, 1954), Rheotanytanytarsus ringei Lehmann
1970, Tanytarsus horni Goetghebuer, 1934b, Tanytarsus mcmillan! Freeman, 1958, Tanytarsus pallidulus
Freeman, 1954, Tanytarsus tri}idus Freeman, 1958, Virgatanytarsus arduennensis (Goetghebuer 1922)
and Virgatanytamts nigricornis (Goetghebuer, 1935).
Examination of additional material from the region has revealed the following new distribution
records: Polypedilum bi/urcatum from Dubai; Cryptochironomus rostratu5 from Syria; Tanytarsus pallidulus
from Yemen; Ablabesmyia longistyla, Chironomus calipterus, Chironomus seychelleanus, Dicrotendipes
freemani, Dicrotendipes sudanicus, Paratendipes nudisquama, Polypedilum nubi/er, Polypedilum
(Tripodura) aegyptium, Stictochironomus puripennis, Cladotanytarsus pseudomancus, Tanytarsus mcmillani,
Tanytarsus tri}idus, Virgatanytarsus arduennensis from Oman; Microchironomus deribae and Polypedilum
nubi/er from Qatar; Paramerina vaillanti Fittkau, 1962, Chironomus calipterus and Polypedilum
(Polypedilum) nubi/erfromJordan and Chironomus calipterusfrom the United Arab Emirates.
From a little further afield, Ablabesmyia longistyla is recorded for the first time from Egypt, Lania
teesdalei from Nigeria, Kie./forulus disparilis from Kenya and the following species from Turkey: Paraphaenocladius
impensus, Paratrichocladius micans, Psectrocladius limbatellus, Dicrotendipes nervosus, Polypeditum
nubeculosum, Polypedilum nubi/er, and Virgatanytarsus arduennensis.
Two as yet unnamed taxa are recorded: Paramerina sp. 1 of LEHMA:-.1N, (1979) and a species of
Chironominae that we cannot allocate to a current genus is briefly described.
ACKNOWLEDGEMENTS
We are grateful to Prof. W. Buttiker for providing Chironomidae from so many localities in
Saudi Arabia and Oman, to my erstwhile colleague Dr. Richard Lane for collections from Oman.
PSC especially thanks Dr. F. Reiss for discussions of problems in taxonomy and biogeography of
south Palaearctic Chironomidae. Annette Seccombe skilfully prepared microscope slides of type
material from the BMNH collections and of material from Turkey and Jordan.
REFERENCES
ADAMS, e. G., 1981. An outline o/Tertiary palaeogeography. In: The Evolving Earth, Cocks, Le., ed.: 221-235. London, British
Museum (Natural History),
ALBlJ, P., 1980. Fam, Chironomidae Sub/am. Chironominae. Fauna Republici Populaire Romine 11 (13): 1-318.
BALL, I. R., 1975, Nature and/ormulation 0/ biogeographic hypotheses, Systematic Zoology 24: 407-430.
BE:>;TON, M.J., 1987. Progress and competition in macroevolution. Biological Review 62: 305-358.
BRUNDlN, L, 1949. Chironomiden und andere Bodentiere der sudschwedischen Urgebirgsseen. Ein Beitrag ZHr Kenntnis der bodenfounistischen
Charakterziige schwedischer oligotropher Seen. Report of the Freshwater Research Institute, Drottningholm
30: 1-915.
BRlJNDI!'.', L., 1966. Transantarctic relationships and their significance, evidenced by chironomid midges. With a monograph ofthe subfomities
Podonominae and Aphroteniinae and the Austral Heptagyiae, Kunglig a Svenska Vetenskapsakadamiens Handlingar
11: 1-427.
CHAlJDHURI, p, K. & D. K. GUHA, 1987. A conspectus 0/chironomid midges (Diptera: Chironomidae) 0/India and Bhutan. Entomol­
ogica Scandinavica Supplement 29: 23-33.
CONTRERAS-LICHTENBERG, R., 1986, Revision der in der Westpaliiarktis verbreiteten Arten des Genus Dicrotendipes Kieffer, 1913
(Diptera, Nematocera, Chironomidae), Annalen des (K. K.) Naturhistorischen (Hof) Museums. Wien 88189: 663-726.
COOPE, G. R., 1979. Late Cenozoic/ossil Coleoptera: Evolution, biogeography and ecology. Annual Review of Ecology and System­
atics, 10: 247-267.
CRACRAIT, J., 1982. Geographic difforentiation, cladistics, and vicariance biogeography: Reconstructing the tempo and mode 0/evolution.
American Zoologist 22: 411-424.

FAUNA OF SAUDI ARABIA 10, 1989
CRANSTON, P. S., 1988. Allergens 0/ non-biting midges (Diptera: Chironomidae): a systematic mrvey 0/ chironomid haemoglobins.
Medical and Veterinary Entomology 2: 117-127.
CRANSTON, P. S., 1989. New species 0/Chironominae (Chironomidae) from Saudi Arabia and the adjacent Middle East. Fauna Saudi
Arabia 10: 225-235.
CRANSTON, P. S. & P. D.ARMITAGE, 1988. The Canary Islands Chironomidae described by T. Becker and E. Santos-Abreu. Deutsche
Entomologische Zeitshrift N.F. 35 (4/5): 341- 354.
CRANSTON, P.S. & P.ASHE, in press. Catalogue of the Chironomidae of the Palaearctic Region. Elsevier.
CRANSTON, P. S., GAD EL RAB, M. O. & A. B. KAY, 1981. Chironomid midges as a cause 0/allergy in the Sudan. Transactions of the
Royal Society of Tropical Medicine and Hygiene 75: 1-4.
CRANSTON, P. S., EDWARD, D. H. D. & D. H. COLLES'>, 1987. Archaeochlus Brundin: a midge out 0/ time (Diptera: Chironomidae).
Systematic Entomology 12: 313-334.
CRANSTON, P. S. & D. R. OLIVER, 1988. Additions and Corrections to the Nearctic Orthocladiinae (Diptera: Chironomidae). Cana­
dian Entomologist 120: 425-462.
CRANSTON, P. S., OLIVER, D. R. & O. A. SAETHER, 1989. The adult males 0/ the Orthocladiinae (Diptera: Chironomidae) 0/ the
Holarctic region - Keys and diagnoses. Entomologica Scandinavica Supplement 32: 164-353.
CRANSTON, P. S., TEE, R. D., CREDL\:'-iD, P. f. & A. B. Kw, 1983. Chironomid haemoglobins: Their detection and role in allergy to
midges in the Sudan and elsewhere. Memoirs of the Entomological Society of America 34: 71-87.
CROIZAT, L., 1958. Panbiogeography. Privately published by the author. Caracas.
CROIZAT, L., NELSON, G. & D. E. ROSEN, 1974. Centres 0/ origin and related concepts. Systematic Zoology 23: 265-287.
DEJOlJX, c., 1984. Contribution to the knowledge o/West African Chironomidae (Diptera-Nematocera). Chironomids/rom the Guinea
Republic. Aquatic Insects 6: 157-167.
DISNEY, R. H. L. & D. H. DE C. HENSHAW, 1988. Be r/es e fluid for slide-mounting insects. Antenna 12: 106-107.
EDWARDS, F.W., 1929. British non-biting midges (Diptera, Chironomidae). Transactions of the Royal Entomological Society of
London 77: 279-430.
EpI.ER, J., 1988. Biosystematics 0/ the genus Dicrotendipes Kie./fel; 1913 (Diptera: Chironomidae) 0/ the world. Memoirs of the
American Entomological Society. 36: 1-214.
FI1TKAU, E.]., 1962. Die Tanypodinae (Diplera, Chironomidae). Die Tribus Anatopyniini, Macropelopiini lind Pentaneurini.
Abhandlung zur Larvalsystematik der Insekten 62: 1-453.
fREEMAN, P., 1954. Chironomidae (Diptera) from (17estern Cape Province - IV Proceedings of the Royal Entomological Society
of London (B) 23: 172-180.
FREEMAN, P., 1955a. Chironomidae (Diptera, Nematocera). Exploration du Parc National Albert. Mission G. F. de Witte
(1933-1935) 83: 3-41.
FREEMAN, P., 1955b. A study 0/ the Chironomidae (Diptera) 0/Africa south 0/the Sahara, Part 1. Bulletin of the British Museum
(Natural History), Entomology 4: 1-67.
FREHI.AN, P., 1956. A study 0/ the Chironomidae (Diptera) 0/Africa south 0/ the Sahara, Part I!. Bulletin of the British Museum
(Natural History), Entomology 4: 285-366.
FREEMAN, P., 1957. A study o/the Chironomidae (Diptera) a/A/rica south o/the Sahara, Part III. Bulletin of the British Museum
(Natural History), Entomology 5: 321-426.
fREEMAN, P., 1958. A study 0/ the Chironomidae (Diptera) o/A/rica south 0/ the Sahara, Part IV Bulletin of the British Museum
(Natural History), Entomology 6: 261-363.
FREEMAN, P., 1961. The Chironomidae (Diptera) 0/Australia. Australian Journal of Zoology 9: 611-737.
FREEMAN, P. & P. S. CRANSTON, 1980. Family Chironomidae. In: Catalogue of the Diptera of the Afrotropical Region. Crosskey,
R.W., ed.: London, British Museum (Natural History). 175-202.
GENTRY, t\. W. & t\.J. SUTCLIFFE, 1981. Pleistocene geography and mammal founas. In: The Evolving Earth. Cocks, L. R. M., (ed.):
London, British Museum (Natural History) 237-251.
GOETGHEBCER, M., 1934a. Ceratopogonidae et Chironomidae du Congo beige (Deuxieme note). Revue de Zoologie et de Bota­
nique Africaine 25: 191-205.
GOETGHEBUER, M., 1934b. Zur Eiforschlmg des persischen Gol/es. (Beitrag Nr. 15). Ceratopogonidae et Chironomidae. Arbeiten
tiber Morphologische u. Taxonomische Entomologie. Berlin-Dahlem 1: 36-39.
GOETGHEBCER, M., 1935. Deux Chironomides du Congo beige. Revue de Zoologie et de Botanique Africaine 26: 397-399.
GOETGHEBUER, M., 1936. Chironomides du Congo beige. Revue de Zoologie et de Botanique Africaine 28: 453-492.
HARE, L. & J. C.II. C>\RTER, 1987. Chironomidae (Diptera, Insecta)/rom the environs 0/a natural West A/rican lake. Entomologica
Scandinavica Supplement 29: 65-74.
HARRISON, A. D., 1971, A conspectus 0/ the Macropelopiini and Pentaneurini (Tanypodinae: Chironomidae) 0/A/rica south 0/ the
Sahara. Canadian Entomologist 103: 386-390.
287

a
288 P. S. CRANSTON, D. D. JUDD
HARRISON, A. D., 1978. New genera and species ofTanypodinae (Diptera: Chironomidae) fiom Afiica south ofthe Sahara. Journal of
the Entomological Society of South Africa 41: 63-80.
HARRISON, A. D., 1987. Chironomidae ofjive central Ethiopian Rift Valley lakes. Entomologica Scandinavica Supplement 29:
39-43.
HIRYENOJA, M., 1973. Revision der Gattung Cricotopus van der Wulp und ihrer Verwandten (Diptera, Chironomidae). Annales
Zoologi Fennici 10: 1-363.
HOLMGREN, A. E., 1869. Bidrag till kannedomen om Beeren Eilands och Spetzbergens insekt-founa. Kungliga Svenska Vetenskaps­
Akademiens Handlingar [4]8(5): 3-55.
HOWARTH, M. K., 1981. Palaeogeography of the Mesozoic. In: The Evolving Earth, Cocks, L. c., (ed.): 197-220. London, Brit­
ish Museum (Natural History).
KIEFFER, J.J., 1908. Chironomidae. Denkschriften Medizinische-Naturwissenschaftlichen Gesellschaft zu Jena 13: 155-162.
KIEFFER, J.J., 1911 a. Description de nouveaux Chironomides de l7ndian Museum de Calcutta. Records of the Indian Museum 6:
113-177.
KIEFFER, J.J., 1911 b. Diptera, Chironomidae der Seychellen-Inseln, aus der Sammlung von Mr H. Scott. Transactions of the Linnean
Society of London, 2nd series, Zoology 14: 331-366.
KIEFFER, J.J., 1912. Description de quatre nouveaux insectes exotique. Bollettino del R. Laboratorio di Zoologia generale e
Agraria della Facolta Agraria in Portici 6: 171-175.
KIEFFER,J.J., 1913. Chironomidae et Cecidomyidae. Resultats Scientifique. Voyage Ch. Allaud et R.Jeannei en Afrique orientale
(1911-1912) (Dipteres). I: 1-43.
KIEFFER, J.J., 1918a. Beschreibung neuer, aufLazarettschifJen des ostlichen Kriegsschauplatzes und bei 19nalino in Litauen von Dr W
Horn gesammelter Chironomiden mit Ubersichtstabellen einiger Gruppen von Palaearktischen Arten (Dipt.). Entomologische
Mitteilungen 7: 35-53; 94-110; 163-170; 177-188.
KIEFFER, J.J., 1918b. Chironomides d"Afiique et Asie conserves au Musee National Hongrois de Budapest. Annales historico-natu­
rales Musei nationalis Hungarici 16: 31-139.
KIEFFER, J.J., 1921 a. Chironomides nouveaux ou peu connus de la region palearctique. Bulletin de la Societe d'Histoire N aturelles
de Metz 29: 51-109.
KIEFFER, J.J., 1921 b. Nouveaux representants de genre Polypedilum (Chironomides). Annales de la Societe Scientifique de Brux­
elles 41: 97-101.
KIEFFER, J.J., 1922. Nouveaux chironomides d larves aquatiques. Annales de Societe scientifique de Bruxelles 41: 355-367.
KIEFFER, J.J., 1923. Chironomides de l'Afiique Equatoriale (3eme partie). Annales de la Societe Entomologique de France 92:
149-204.
KIEFFER, J.J., 1924. Six nouveaux Chironomides d"Afiique. Annales de la Societe Scientifique de Bruxelles 43: 255-261.
KIEFFER, J.J., 1925. Chironomides d'Egypte (Dipt.). Bulletin de la Societe Entomologique d'Egypte 8: 244-313.
KITCHING, 1.J., VANE-WRIGHT, R.1. & P. R.AcKERY, 1987. The Cladistics ofIdeas. Cladistics 3: 14-34.
KUGLER, J., 1966. Vorliiujige Mitteilung iiber die Chironomiden-Fauna des Tiberiassees. Gewasser und Abwasser 41/42: 70-84.
KUGLER, J. & F. REISS., 1973. Die triangularis-Gruppe der Gattung Tanytarsus v.d. W (Chironomidae, Diptera). Entomologisk
Tidskrift 94: 59-82.
KUGLER, J. & WOOL, 1968. Chironomidae (Diptera)fiom the Hula Nature Preserve, Palestine. Annales Zoologi Fennici 5: 76-83.
LANGTON, P. H., 1980. The genus Psectrocladius Kieffir (Diptera: Chironomidae) in Britain. Entomologist's Gazette 31: 75-88.
LANGTON, P. H., 1984. Review oftype species of the limbatellus group, with a provisional key to known fimales ofPsectrocladius
Kieffir (Diptera: Chironomidae). Entomologica Scandinavica 15: 477-485.
LARSEN, T. B., 1984. The zoogeographic composition and distribution ofthe Arabian butterflies (Lepidoptera: Rhopalocera). Journal
of Biogeography 11: 119-158.
LEHMANN, J., 1970. Revision der europiiischen Arten (Imagines d'd' und Puppen d'd') der Gattung Rheotanytarsus Bause (Dipt.,
Chironomidae). Zoologische Anzeiger 185: 344-378.
LEHMANN, J., 1979. Chironomidae (Diptera) aus Fliessgewassern Zentralafiikas. TeilI: Kivu-Gebiet, Ostzaire. Spixiana Supplement
3: 1-144.
LEHMANN, J., 1981. Chironomidae (Diptera) aus Fliessgewassern Zentralafiikas. Teil II: Die Region urn Kisangami, Zentralzaire.
Spixiana Supplement 5: 1-85.
LINNAEUS, c., 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum caracteribus, differentiis,
synonymis, locis. Ed. 10. 1, 824 pp. Holmiae (= Stockholm).
MEIGEN, J.J., 1804. Klassification und Beschreibung der europaischen zweiflUglichen Insekten (Diptera Linn.). I. Erste Abtheilung,
XVIII + 1-152; zweite Abtheilung, VI + 153-314. Braunschweig (= Brunswick), Reichard.
MEIGEN, J.J., 1818. Systematische Beschreibung der bekannten europaischen, zweifliigeligen Insekten. 1, XXXVI + 332 pp.
Aachen, Forstmann.