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A revision of Afrotropical Quasimodo flies (Diptera: Schizophora; Curtonotidae). Part III--the Malagasy species of Curtonotum Macquart, with descriptions of six new species.

INTRODUCTION

Four genera of Curtonotidae are currently known worldwide, i.e., Axinota van der Wulp, 1886, Cyrtona s.l. Seguy, 1938, Tigrisomyia Kirk-Spriggs, 2010 and Curtonotum Macquart, 1844, all of which occur in the continental Afrotropical Region; Tigrisomyia exclusively so. Curtonotum is the most speciose genus and is found in all zoogeographical regions of the world except Australasia/Oceania and Antarctica, although Klymko and Marshall (2011) point out that Curtonotum in its current broad sense is paraphyletic with respect to Axinota, and suggest restricting the name Curtonotum to a monophyletic New World group. Two genera are known to occur on Madagascar, Axinota, represented by a single species, A. kyphosis Kirk-Spriggs, 2010, of supposed Oriental origin and Curtonotum (Appendix I). Notably, the genus Cyrtona s.l., which is extremely species-rich in the continental Afrotropical Region, is apparently absent.

The Curtonotum fauna of Madagascar has remained a neglected group and nothing is currently known of their biology and immature stages. Species in the continental Afrotropical Region are known to roost in the burrows of small mammals, hollow trees and overhangs of riverbeds, etc. (e.g., Kirk-Spriggs 2008b; Meier et al. 1997; Tsacas 1977) and, at least in the case of species occurring in xeric regions, are known to develop as scavengers in the damaged egg pods of locusts and grasshoppers (Orthoptera) (Greathead 1958; Kirk-Spriggs 2008b; Meier et al. 1997). A more complete review of the known biology of the genus will be presented in Part IV of this revision.

Specimens of C. balachowskyi Tsacas from Madagascar are subject to infestations of the entomophagous fungus Laboulbenia curtonoti Rossi & Kirk-Spriggs (Ascomycota), a species with an elongated rhizoid which penetrates the host's cuticle (Rossi & Kirk-Spriggs 2011). An infestation of a different, undescribed Laboulbeniales species was also observed on a specimen of C. sakalava Tsacas during the course of this study, but occurred in insufficient numbers to facilitate description.

In a first attempt to deal with the taxonomy of the Madagascan species of the genus, Tsacas (1974) described and named seven endemic species and identified, but did not name, five apparent others. Tsacas' study was, however, based on only 38 specimens. These were collected by Renaud Paulian (1913-2003), then Deputy Director of the Institut Scientifique de Madagascar, and colleagues from 1956 to 1960. This was supplemented with material sampled by Raymond Decary (1891-1973) in 1922, and Alfred "Fred" Jakob Keiser-Jenny (1895-1969) and his wife collected from May to October 1958. The study also included specimens collected by Brian Roy Stuckenberg (1930-2009), during his second Madagascan expedition from November 1957 to April 1958.

All specimens studied by Tsacas (1974), including all types, were borrowed for study from the respective institutions. A few additional specimens came to light in material loaned from Tel Aviv University, Israel, Zoologische Staatssammlung, Munchen, Germany, and the KwaZulu-Natal Museum, Pietermaritzburg, South Africa, but by far the most substantial number of specimens now available for study result from Michael E. Irwin's extensive Arthropod Survey of Madagascar's Protected Areas (1998-2009), deposited in the California Academy of Sciences, San Francisco, USA.

Tsacas (1974) illustrated the spermathecae of five named and two unnamed Madagascan species, but did not describe these structures or assess their variability. Kirk-Spriggs (2008a), however, assessed intra- and inter-specific variability in spermathecal form of three continental Afrotropical species in the Curtonotum cuthbertsoni complex (sensu Tsacas 1977), with obclavate spermathecae and concluded that variability was too great to allow specific differentiation. For this reason (with the notable exceptions of C. sternithrix, which has dark rings around the sockets of all thoracic setae, and C. sakalava, which is unique in having the lateral maculae of the abdominal tergites developed into a continuous band in both sexes), only the males can be identified with certainty and then only following detailed examination of the male terminalia.

As noted above, Tsacas (1974) identified, but did not name, five "species" from Madagascar (Curtonotum sp. cf. balachowskyi n. sp. species a, C. cf. balachowskyi n. sp. species b, C. sp. cf. pauliani, n. sp. species c, Curtonotum sp. d, and Curtonotum sp. e), only the first of which is represented by a male and can be reliably determined. Close examination of the male terminalia of this specimen, in direct comparison to the types, reveals its conspecificity with C. balachowskyi. As these specimens were not formally named they have no bearing in nomenclature and the remaining female specimens must be regarded as indeterminate Curtonotum spp.

Earlier parts of this revision deal with the genus Axinota (Kirk-Spriggs 2010a) and the East African endemic genus Tigrisomyia (Kirk-Spriggs 2010b). Part IV will consider the species of Curtonotum occurring in the continental part of the Afrotropical Region and will include a more thorough account of species complexes and group associations.

MATERIAL AND METHODS

Preparation of spirit-preserved specimens

Material resulting from the Arthropod Survey of Madagascar's Protected Areas (1998-2009) and other material loaned from the California Academy of Sciences was sampled using Malaise traps and was initially preserved in 96 % ethanol. Only part of the material resulting from this extensive survey was available for study as numerous samples remain unsorted. These specimens were prepared in the following manner: specimens were placed on dry tissue paper to absorb excess alcohol and were then placed in vials of 2-ethoxy ethanol and left to stand overnight. The 2-ethoxy ethanol was then decanted and replaced with ethyl acetate, and again left overnight. Specimens were then removed and air-dried on tissue paper before being glued to card points using Seccotine and having the appropriate labels added. Ex-alcohol specimens are bleached to some extent and many have been rubbed or have setation missing; colours of dry-pinned specimens can be expected to be brighter.

Dissection

All male specimens included in this study were dissected. Abdomens were removed with watchmaker's forceps and macerated in 10 %<> potassium hydroxide in a heated block before being transferred to 70 % ethanol with a few drops of glacial acetic acid in an excavated glass block. Following dissection, abdomens and male terminalia were stored in a micro-vial with glycerol pinned beneath each specimen. The cork-bunged glass genitalia vials used by L. Tsacas, many of which had dried up or deteriorated, were all replaced and a label was appended to each specimen stating "Genitalia vial replaced A.H. Kirk-Spriggs [2006-2008]".

Preparation of figures and images

Figures of the head, thorax and frons (Figs 1-26), were prepared from images captured with a [R]Leica EZ4HD stereo microscope with built-in digital camera. These were photographed using a ring light and dome (Kerr et al. 2008) in a range of focal planes and were digitized using [R]Combine ZP Image Stacking Software.

Wings were detached and placed between two glass slides. Images (Figs 27-35, 3739) were captured with a [R]Leica Wild M3Z binocular microscope with [R]Nikon E5400 digital camera attached. Fig. 36 was prepared using the [R]Leica EZ4HD microscope.

Male terminalia were mounted laterally or dorsally, in a blob of heated and then cooled glycerine jelly. Figures of these structures (Figs 40-91) were prepared using a [R]Leica Wild M3Z binocular microscope with a camera lucida attachment. Images were enlarged and traced, and details added by hand in pen and ink by constant referral to the specimen. Scale bars were added using a 0.5 mm graticule slide.

Measurements

Measurements were taken with a graticule eye piece on a [R]Leica Wild M3Z binocular microscope, calibrated with a 0.5 mm graticule slide.

As the descriptions and redescriptions are based on the holotype in most cases, measurements are provided for the holotype only. In some cases overall lengths of series of specimens were taken prior to dissection and in such cases a range of lengths is provided. In some cases overall lengths could not be provided as specimens were dissected beforehand. In such cases the measurements provided by Tsacas (1974) are included (where applicable), or combined length of the head and thorax and thorax and scutellum alone are provided. Tsacas did not specify how his measurements were made, but overall length measurements appear to have included the wings (i.e., measured from the head to the tips of the wings), rather than the combined lengths of the head, thorax and abdomen, so are larger by comparison with the measurements provided here.

Overall length was measured from the anteriormost point of the frons to the tip of the abdomen (viewed laterally). Length of head and thorax combined was measured from the anteriormost point of the frons to the scutoscutellar suture (viewed dorsally).

Length of thorax and scutellum combined was measured from the anterior margin of the mesonotum (immediately posterior to head) to the anterior tip of the scutellum (viewed dorsally).

The eye height/length ratio was measured from the most ventral to the most dorsal point of the eye and through the eye medially (viewed laterally).

Genal height was measured immediately below the ventromedial part of the eye (viewed laterally).

Frons length represents the distance between the ventral margin (immediately above the point of antennal insertion), to the posterior ocelli (viewed dorsally). Frons width represents the distance between the lateral margins of the frons, measured at the midpoint between the posterior ocelli and the ventral margin of the frons.

Length of the wing was measured from the humeral crossvein (Fig. 27, h) to the apex of the second radial vein (Fig. 27, [R.sub.4+5]).

Descriptions

A full redescription is provided for C. balachowskyi Tsacas, the most widely distributed species of the genus in Madagascar. Redescriptions and descriptions of other species are compared to this and differences and additional characters noted only. A full description or redescription of the male terminalia is provided in all cases.

Distribution maps

Each set of specimen locality co-ordinates listed in Appendix II was obtained in one of two ways: (a) original co-ordinates provided on data labels, later converted to decimal co-ordinates; or (b) obtained from Internet geo-referencing websites powered by Google Earth(tm).

Distribution maps were prepared by converting the list of geographic co-ordinates indicating the point location of specimen records into a spatial data layer in ArcMap 9.3. The specimen location layer was overlaid on the Vegetation Map of Madagascar by Moat and Smith (2007). Plotted points are based on material examined as part of the systematic revision only, i.e., all represent confirmed identifications.

Vegetation types cited in the text follow Moat and Smith (2007), as listed in Appendix II, biomes follow Yoder and Nowak (2006) (Fig. 105), biogeographical zones Boumans et al. (2007) and Wilme et al. (2006) (Fig. 106) and bioclimatic zones Cornet (1974) and Schatz (2000) (Fig. 107).

Labels

Type label data are quoted exactly as they appear. A division slash (/) denotes the commencement of a new line, two division slashes (//) data on a further label. Significant supplementary or qualifying information is presented in square brackets when considered necessary. Information on label colour, etc. is only provided for type material. All labels are printed on white card unless otherwise stated. Specimens with the head missing are labelled as such, e.g., "Head missing 2008". Adult morphology follows Cumming and Wood (2009) and for the male terminalia Marshall et al. (2010).

Institutional codens

A list of institutional codens used in the text is provided below, with the names of respective curators and collection managers noted in parenthesis.

BMSA--National Museum, Bloemfontein, South Africa.

CAS--California Academy of Sciences, San Francisco, USA (N. Penny, C. Griswold). CAS specimens have collection codes with the prefix 'MA-' or 'MG-'.

FBUB--Biological Collection, Universitat Bielefeld, Bielefeld, Germany (M. von Tschirnhaus).

MNHN--Museum national d'Histoire naturelle, Paris, France (Ch. Daugeron).

NHMB--Naturhistorisches Museum, Basel, Switzerland (D. Burckhardt, I. Zucher).

NMSA--KwaZulu-Natal Museum, Pietermaritzburg, South Africa (M. Mostovski, R. Miller, P. Birkett).

TAU--Tel Aviv University, Tel Aviv, Israel (A. Freidberg).

ZSM--Zoologische Staatssammlung, Munchen [= Munich], Germany (M. Kotrba, B. Stock, some via Michael von Tschirnhaus (FBUB)).

Abbreviations used in the text, on figures and in figure legends

[A.sub.1] + Cu[A.sub.2]--first anal vein (fold); [A.sub.2]--second anal vein (fold); AT--allotype; bp--basiphallus; C--costa; ce--cercus; [cua.sub.1]--cubital cell; CuA1--cubital vein; dm--discal medial cell; dm-cu--discal medial-cubital crossvein; dp--distiphallus; ea--ejaculatory apodeme; ep--epandrium; h--humeral crossvein; HT--holotype; hy--hypandrium; m--medial cell; M--medial vein; m--metres above sea level; N-T--non-type/s; pg--postgonite; ph--phallapodeme; PT--paratype/s; r1--first radial cell; [r.sub.2+3]--second radial cell; [R.sub.2+3]--second radial vein; [r.sub.4+5]--third radial cell; [R.sub.4+5]--third radial vein; r-m--radial-medial crossvein; Sc--subcosta; ss--surstylus.

TAXONOMY

Genus Curtonotum Macquart, 1844

Refer to Appendix I for synonymy.
Key to males of Malagasy species of Curtonotum

(Examination of the male terminalia is required in all cases.)

1 Wing membrane deep brown infuscate throughout (Figs 27-29);
  distiphallus either with narrow, ventrally-directed, lateral spine
  and two smaller basal spines (Figs 61, 62, indicated with arrows),
  or with patch of finger-like spinules basally and tuft
  of hairs submedially (Fig. 60)                                      2

--Wing membrane only faintly brown infuscate, sometimes marginally
  darker in r1 and anterior half of r2+3 and/or in region of dm-cu
  crossvein only (Figs 30-39); distiphallus lacking
  ventrally-directed spines or other spinular modifications (if
  spines present, then these forming anterior margin of basiphallus)  4

2 Three strong postpronotal setae; all setae on mesonotum and
  thoracic pleura with distinct dark brown ring around each socket
  (Fig. 3); extensively developed lateral maculae and medial fascia
  of abdominal tergites black; sternites 4 and 5 modified, with comb
  of long, regular spinules on anterior margin, sternite 6 with deep
  V-shaped apical excision (Fig. 86); distiphallus with patch of
  finger-like spinules basally and tuft of hairs submedially (Fig.
  60)                                                sternithrix Tsacas

--Two strong postpronotal setae; all setae on mesonotum and
  thoracic pleura without distinct dark brown ring around each socket
  (Figs 1, 2); extensively developed lateral maculae and medial
  fascia of abdominal tergites brown; sternites 4 and 5 unmodified,
  sternite 6 dove-tailed (Figs 79, 80); distiphallus with
  conspicuous, ventrally-directed, long, lateral spine and two
  smaller spines (Figs 61, 62, indicated with arrows)                 3

3 Frons (Fig. 14) pale brown, with conspicuous deep brown vittae
  reaching ventral margin; dm-cu crossvein angle and curvature as
  illustrated (Fig. 27); phallus weakly sclerotised at lateral
  margins of basiphallus (viewed dorsally) (Fig. 61); prominent
  ventrally-directed spine on distiphallus narrow, smaller spines
  positioned in basolateral region (Fig. 61)             keiseri Tsacas

--Frons (Fig. 15) pale yellow, slightly darker posteriorly and
  medial to orbital plates (if frons slightly darker, vittae never
  strong or reaching ventral margin); dm-cu crossvein angle and
  curvature as illustrated (Fig. 28); phallus heavily-sclerotised at
  lateral margins of basiphallus (viewed dorsally) (Fig. 62);
  prominent ventrally-directed spine on distiphallus broad, smaller
  spines positioned in left and right lateral regions (Fig. 62)
                                                    stuckenbergi Tsacas

4 Epandrium (viewed laterally) with oblique semicircular excavation
  along ventroapical margin, ventral lobe forming a blunt point, from
  which two very strong, long black setae originate (Figs 46, 47);
  anterior margin of basiphallus with finger- or spine-like
  ventromedial projection (Figs 49, 50, 52, 53)                       5

--Epandrium (viewed laterally) evenly-rounded or straight at
  ventroapical margin, with extensive group of irregular, long setae
  along ventral margin (Figs 40, 41, 48, 64-66, 73, 76); anterior
  margin of basiphallus without finger- or spine-like ventromedial
  projection                                                          6

5 Frons pale dirty yellow, faintly darker towards vertex and
  between orbital plates, both vittae inconspicuous (Fig. 23);
  sternite 6 with dense setulae (Fig. 88); surstylus short, wide
  basally (Fig. 46); dm-cu crossvein angle and curvature as
  illustrated (Fig. 36); basiphallus evenly rounded basally, with
  conspicuous internal bulge in basal fifth (Fig. 49); distiphallus
  with medial finger-like, slightly curved, ventrally-directed
  projection (Figs 49, 52)                                 boeny Tsacas

--Frons pale brown to dark brown, forming conspicuous vittae,
  reaching ventral margin (Fig. 24); sternite 6 with sparse setulae
  (Fig. 89); surstylus long, narrow basally (Fig. 47); dm-cu
  crossvein angle and curvature as illustrated (Fig. 37); basiphallus
  straight in basal third, with slight internal bulge at midlength
  (Fig. 50); distiphallus with medial spine-like, ventrally-directed
  projection (Figs 50, 53)                             griveaudi sp. n.

6 Phallus highly modified; basiphallus (Fig. 74) grossly expanded,
  forming extensive, extremely wide, sclerotised, half moon-shaped
  plate, strongly concave on left side, with large, heavily
  sclerotised sperm pump (Figs 74a, 74b), possibly inserted in left
  concavity; dm-cu crossvein angle and curvature as in Fig. 35
                                                     rinhatinana sp. n.

--Phallus not highly modified; basiphallus not grossly expanded,
  narrow (Figs 42, 43, 51, 67-69, 77); sperm pump absent; dm-cu
  crossvein angle and curvature not as in Fig. 35)                    7

7 Hypandrium short, with broad-based subangulate-truncate
  dorsobasal lobe (Figs 48, 76); sternite 6 as illustrated (Figs 90,
  91); basal section of distiphallus either with a conspicuous,
  short, dorsomedial keel (Figs 77, 78), or with conspicuous, short
  dorsomedial spine (Figs 51, 54)                                     8

--Hypandrium long, with broad-based rounded or subtriangular dorsobasal
  lobe (Figs 40, 41, 64-66); sternite 6 as illustrated (Figs 81-85);
  distiphallus lacking a keel or short, dorsomedial spine             9

8 Very small species, wing length ca 1.6 mm; frons (Fig. 26) markedly
  wider at vertex than at ventral margin; ocellar triangle large,
  extending ca one third length of frons; lateral margins of tergites
  2-4 collectively forming an unbroken facia (at low magnification);
  dm-cu crossvein angle and curvature as in Fig. 39; distiphallus with
  conspicuous, short, dorsomedial keel (Figs 77, 78)
                                                        sakalava Tsacas

--Larger species, wing length ca 3.2 mm; frons (Fig. 25)
  subparallel-sided; ocellar triangle small, extending ca one-fifth
  length of frons; lateral margins of tergites 2-4 with well-separated,
  subelliptical macula basally; dm-cu crossvein angle and curvature as
  in Fig. 38; distiphallus with short, dorsomedial spine (Figs 51, 54)
  (Madagascar, Namibia and South Africa)                pauliani Tsacas

9 Sternite 6 broader than long, subquadrate (Figs 84, 85), either with
  very short setulae, only marginally longer at apical margin (Fig.
  84), or with extremely long setulae at apical margin (Fig. 85);
  basiphallus in apical region, either with distinct raised, angulate
  fold on left margin (Fig. 44), or with two spines, one medial and
  one submedial (Fig. 45)                                            10

--Sternite 6 narrower in basal ca third, expanded laterally, with
  V-shaped or Ushaped apical excision, with moderately strong setulae
  at apical margin (Figs 8183); basiphallus (in apical region) either
  with left lateral margin developed into a long, curved, ventrally-
  directed spine and smaller medial spine (Fig. 70), or with distinct
  raised keel and spinose area laterally on distiphallus (Figs 71,
  72)                                                                11

10 Frons (Fig. 20) subparallel-sided, pale dirty yellow, faintly darker
   towards vertex and between orbital plates, otherwise two vittae
   inconspicuous; dm-cu crossvein angle and curvature as illustrated
   (Fig. 33); basiphallus (Fig. 44) with right lateral margin evenly
   rounded, with slightly raised margin, forming even projection
   apically, left lateral margin with distinct, angulate, raised fold
                                                    balachowskyi Tsacas

--Frons (Fig. 21) markedly wider at vertex than at ventral margin, pale
  brown, darker brown towards vertex and between orbital plates,
  forming conspicuous vittae that reach ventral margin; dm-cu
  crossvein angle and curvature as illustrated (Fig. 34); basiphallus
  (Fig. 45) with right lateral margin evenly rounded and flat,
  developed into a bluntly pointed ventromedial spine, with much
  smaller spine basolaterally                       gladiiformis sp. n.

11 Basiphallus moderately narrow in apical third (viewed dorsally),
   with left lateral margin developed into a long, curved,
   ventrally-directed spine and smaller medial spine (Fig. 70); dm-cu
   crossvein angle and curvature as illustrated (Fig. 30) irwini sp. n.

--Basiphallus laterally expanded in apical third (viewed dorsally)
  (Figs 71, 72), with distinct raised keel in left apical region and
  spinose area on lateral margin of distiphallus; dm-cu crossvein
  angle and curvature not as Fig. 30                                 12

12 Distiphallus (Fig. 71) with sclerotised area subdivided medially,
   with thin ventrally directed spine in membranous window, right
   lateral section with three short, darktipped spines (viewed
   dorsolaterally; not visible on Fig. 71); dm-cu crossvein angle and
   curvature as illustrated (Fig. 31)                    parkeri sp. n.

--Distiphallus (Fig. 72) with basolateral, upturned, sclerotised,
  crown-like projection, this subtriangular (viewed laterally) (Fig.
  69); dm-cu crossvein angle and curvature as illustrated (Fig. 32)
                                                   coronaeformis sp. n.


Curtonotum balachowskyi Tsacas, 1974

Figs 7, 20, 33, 40, 42, 44, 84, 98

Curtonotum balachowskyi: Tsacas 1974: 710; figs 4a-d (p. 711); fig. 8f (p. 718). Type locality: "Madagascar: Mananjary".

Curtonotum sp. cf. balachowskyi n. sp. species b (sensu Tsacas 1974: 712).

Differential diagnosis: This species differs from other species occurring on Madagascar in having the medial lobes of the hypandrium (viewed dorsally), parallel-sided, with a narrow medial membranous area clothed in tiny spinules. Interpretaton of its relationships to other species occurring in the Afrotropical Region must await the outsome of phylogenetic study.

Redescription:

Male (primarily based on field-pinned HT).

Measurements: Overall length 2.8-4.8 mm (n = 47, N-T); length of head and thorax combined 2.8 mm; length of thorax and scutellum combined 2.9 mm; wing 4 mm long. Head (Figs 7, 20). Compound eye probably green-brown iridescent in living examples, in profile gently and evenly rounded anteriorly, slightly triangularly produced posteriorly, eye height/length ratio: 12:7 (HT); frons (Fig. 20) subparallel-sided, slightly wider than long, frons length/width ratio: 7:85 (HT), very slightly wider at vertex than at ventral margin, ground colour pale dirty yellow, faintly darker towards vertex and between orbital plates, otherwise both vittae inconspicuous, surface with a few minute pale to dark brown setulae, positioned at medial margin of orbital plates; orbital plates and ocellar triangle silver-grey pruinose; ocelli clear grey with 4 or 5 minute dark setulae arranged in two closely-approximated regular rows between posterior ocelli; orbital plates extending from vertex of head to 0.9 length of frons, slightly indented at lateral margin between posterior and anterior orbital setae; lateral margins with narrow silver pruinose fascia (adjacent to eye margin), widest at antennal insertions; posterior orbital seta moderately strong, slightly outcurved, shorter than outer vertical seta, with tiny proclinate medial orbital seta inserted anteromedially to socket of posterior orbital seta; anterior orbital seta moderately strong, ca half length of ocellar setae; ocellar setae finer, almost reaching ventral margin of frons; outer vertical seta shorter than inner; postocellar setae strong, cruciate, slightly shorter than outer vertical seta; antennal scape and pedicel dirty pale brown, silver-grey pruinose, flagellomere 1 concolourous with pedicel basally, darkened apically, silver-grey pruinose as in face, longer than wide, apex bluntly-pointed, arista with 9-11 long dorsal branches and 3 or 4 ventral branches in addition to terminal fork; lunule and face uniform silver-grey pruinose throughout, face with broad silver fascia (adjacent to eye margin), facial carina developed as a low ridge, extending half length of face, pre-epistomal line indented; clypeus brown, especially laterally; 1 pair fairly strong vibrissae inserted on posterior lateral margin and 10 much finer setae bordering genal groove; occiput yellow to grey pruinose with moderately strong, black postocular setae; gena narrow, eye height/genal height ratio: 12:1 (HT), silver pruinose, abruptly dirty brown beyond basal angle; palpus black-brown, brown microtrichose.

[FIGURES 1-8 OMITTED]

Thorax (Fig. 7). Mesonotum moderately convex, very slightly flattened behind head, with multiple rows of regular, short, black, overlapping setulae; silver-grey pruinose, with four parallel chestnut-brown pruinose vittae on dorsal surface, 2 median vittae extending from anterior margin to region of anterior dorsocentral seta socket, 2 lateral vittae shorter, extending from % length to region of posterior dorsocentral seta socket, 2 pairs of dorsocentral setae, posterior long and strong (shorter than lateral scutellar seta), anterior shorter and finer (shorter and finer than medial scutellar seta); 1 pair acrostichal setae shorter than anterior dorsocentral seta; presutural seta moderately strong, reclinate, as long and strong as posterior notopleural seta; 2 notopleural setae the anterior slightly longer than posterior; 1 strong, reclinate supra-alar seta slightly exceeding length of posterior dorsocentral seta; 2 reclinate postalar seta moderately strong, same size as acrostichal setae; postpronotum dirty yellow-grey pruinose, with 2 strong postpronotal setae, the more dorsal longer and reclinate, the more ventral shorter and proclinate, with 10-12 finer black-brown setulae; anepisternum silver-grey pruinose with 3 moderately strong anepisternal setae, the dorsal and medial reclinate, stronger and of equal length, the more ventral shorter, slightly dorsally-directed, surface with 18 fine setulae scattered across surface, some larger and arranged in two groups of 3; anepimeron, laterotergite and meron silver-grey pruinose, glabrous; katepisternum silver-grey pruinose, with 2 katepisternal setae, the more ventral strong, slightly dorsally-directed, the more dorsal much smaller and finer, ca 0.2 length of ventral katepisternal setae, surface with 16 short, fine setulae at base and along posterior margin.

Scutellum. Silver-grey pruinose as in mesonotum, with faint medial brown pruinose vitta basally (under some lights); anterior 0.8 clothed in black, irregular, overlapping setulae; 2 pairs of strong scutellar setae, 1 weak basal scutellar setula and 1 weak intermediate scutellar setula, the latter inserted closer to lateral than medial scutellar setae (0.8 distance between medial and lateral scutellar setae).

Legs. Fore coxa silver-yellow pruinose with two moderately strong, brown, ventrally-directed preapical setae and comb of finer setulae medially, with 8-10 brown setulae on anterior surface; mid and hind coxa yellow-grey pruinose, mid coxa with 2 very strong, lateral, ventrally-directed, black setae and comb of finer setae medially and 3 brown setulae; hind coxa with 1 weaker lateral black seta and 1 brown setula; femora, tibia and tarsi uniform dirty yellow; all tibiae with preapical seta; fore tibia with 4 strong setae on lateral margin, the second basal seta shorter than other three, with ctenidium of 10-12 short, sharp, black spinules, separated from each other by one or more basal spinule widths.

Wing (Fig. 33). Long, relatively narrow, tip evenly-rounded, veins chestnut-brown, membrane very faintly infuscate brown throughout, very slightly darker in [r.sub.1] and anterior half of [r.sub.2+3] and in region of dm-cu crossvein; costa with prominent costal spines in basal 0.8 from costal break; [R.sub.4+5] slightly anteriorly-produced in basal third, [R.sub.2+3] and [R.sub.4+5] subparallel, [r.sub.2+3] expanded apically; dm-cu crossvein slightly obliquely angled posteriorly, evenly curved medially; [cua.sub.1] relatively long and narrow; [A.sub.1]+Cu[A.sub.2] and [A.sub.2] manifested as a fold only; [A.sub.2] extending half length of cuap haltere dirty yellow.

Abdomen. Ground colour of tergites 1-5 yellow-grey pruinose, clothed in relatively long black, overlapping setulae, arranged in regular rows, those at apical margins longer and stronger; tergite 1 simple, devoid of maculae; tergite 2 with oblique, sub-rectangular brown-black pruinose dorsolateral macula on either side only; tergites 3-5 with narrow, V-shaped concolourous median fascia and well-separated and reduced concolourous T-shaped dorsolateral macula, lateral margin of tergites 2-5 with subelliptical concolourous macula in basal half; sternite 4, quadrate, sternite 5 rectangular, subparallel-sided, 0.25X longer than sternite 4, both unmodified, with brown setulae arranged in irregular rows, those along apical margins slightly longer and stronger; sternite 6 (Fig. 84) subquadrate (may appear narrower in undissected specimens), narrowed in basal 0.4, with shallow, broad, subtriangular excision apically, with faint brown maculae medially and fascia laterally, merging apically, clothed in short black irregular brown setulae in apical 2/3, those at apical margin longer and more prominent.

Terminalia (Figs 40, 42, 44). Hypandrium (Fig. 40, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (rounded to slightly angulate in profile); hypandrial arms narrow basally, expanded apically (viewed laterally), with 2 setulae proximal to postgonite, the more lateral ventrally directed, the medial ventromedially directed (obscured by epandrium on Fig. 40), sclerotised area of medial lobes (viewed dorsally), parallel-sided, medial membranous area narrow, clothed in tiny spinules, separated in basal half, slightly overlapping in apical half; postgonite (Fig. 40, pg) long, thin and straight, with slight undulating anterior margin; epandrium (Fig. 40, ep) slightly broader dorsally than ventrally (viewed laterally), evenly-rounded on dorsal margin, posterior margin slightly angled, ventral margin with extensive row of long, regular to irregular, apically-directed setae; cercus (Fig. 40, ce) not prominent, longest setae as long as medium setae on ventral margin of epandrium; surstylus (Fig. 40, ss) long, widest basally, slightly curved in apical %; phallus (as in Figs 42, ph, bp, dp; 44, bp, dp) C-shaped, moderately sclerotised, brown; phallapodeme (Fig. 42, ph), fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two short, flat projections in basal 0.4, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 42, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 42, bp) narrow and regular for most of its length, with very slight internal bulge at point of first bend in basal three-fifths; apical section (Fig. 44, bp) broad, right lateral margin evenly rounded with slightly raised margin, forming even projection apically, left lateral margin with distinct raised fold; distiphallus (Figs 42, 44, dp) long, scimitar-like (viewed laterally), narrow (viewed dorsally).

Variation: The even projection on the right anterior margin of the basiphallus is somewhat variable in shape and angle in some specimens, other terminalia characters are consistent, however, and this is here regarded as intraspecific variation only.

Holotype (examined): [male] MADAGASCAR: "MADAGASCAR Fia / Mananjary / 11.VIII.[19]58.F.KEISER [pink paper] // HOLOTYPE [red card] // CURTONOTUM / balachowskyi / [male] holotype n.sp. / L. TSACAS DET. 1974 [printed & handwritten] // MUSEUM PARIS // Curtonotum / balachowskyi [male] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit 2008" (MNHN). In excellent condition; re-staged on new poly mount; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "Curtonotum/ balachowskyi [male] [or [female]] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit2008"): MADAGASCAR: 1[female] same data as holotype except: "// PARATYPE [printed; red card] // CURTONOTUM / balachowskyi n.sp. / $ paratype / L. TSACAS DET. 1974 [printed & handwritten]"; 1[female] same except: "// CURTONOTUM / balachowskyi n.sp. / [female] paratype / L. TSACAS DET. 1974 [printed & handwritten]" (both NHMB); 1[female] same except: "6.VIII.[19]58 // ALLOTYPE [printed; red card] // CURTONOTUM / balachowskyi / [female] allotype n.sp. / L. TSACAS DET. 1974 [printed & handwritten]"; 1[male], "Ambila / [?Sunaibo--illegible] [handwritten] // VIII.[19]52 / (R.P.) [= R. Paulian] [handwritten] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [pale grey card] // PARATYPE [red card] // CURTONOTUM / balachowskyi / [male] paratype n.sp. / L. TSACAS DET. 1974 [printed & handwritten] // MUSEUM PARIS" [head detached, glued to data label] (both MNHN); 1[male], "MADAGASCAR Tam. / Tamatave / 23.X.[19]58 F.KEISER [pink paper] // PARATYPE [red card] // CURTONOTUM / balachowskyi / [male] paratype n.sp. / L. TSACAS DET. 1974 [printed & handwritten]" (NHMB).

Other material examined (all labelled: "Curtonotum balachowskyi Tsacas, 1974 [male] det. A.H. Kirk-Spriggs 2010"): MADAGASCAR: d'AnalalavaProv.: 1[male] Maromandia, R. Decary 1922, "Curtonotum sp. cf. balachowskyi n. sp. espece b, L. Tsacas det. 1974" (MNHN): Fianarantsoa Prov.: 1[male] near Isalo National Park, at stream east of Interpretive Center, 22[degrees]37.60'S:45[degrees]21.49'E, 13-19.i.2002, 750 m, M.E. Irwin & R. Harin'Hala, Malaise trap in open area (MA-02-11A-11); 1[male] near Isalo National Park, in dry wash east of Interpretive Center, 22[degrees]37.60'S:45[degrees]21.49'E, 12-22.vi.2002, 885 m, M.E. Irwin & R. Harin'Hala, Malaise trap in open area (MA-02-11B-32); 4[male] 50 km S of Farafangana, Mahabo Mananivo, Ampitavananima Forest, 23[degrees]7.79'S:47[degrees]43.02'E, 13-20.i. 2007, 34 m, M.E. Irwin, F. Parker & R. Harin'Hala, Malaise trap low altitude littoral rainforest (MG-35-01); same except: 1[male] 20-26.i.2007 (MG-35-02); 1[male] 26.i-4.ii.2007 (MG-35-03); 1[male] 4-10.ii.2007 (MG-35-04); 1[male] 10-17.ii.2007 (MG-35-05); 1[male] 17-24.iii.2007 (MG-35-10); 1[male] 24-31.iii.2007 (MG-35-11); 2[male] 14-21.v.2007 (MG-35-18); 5[male] 21-25.v.2007 (MG-35-19); 1[male] 29.v-2.vi.2007 (MG-35-20); 1[male] 2-10.vi.2007 (MG-35-21); 1[male] 10-15.vi.2007 (MG-35-22); 2[male] 15.vi-1. vii.2007 (MG-35-23); 5[male] 14-19.vii.2007 (MG-35-26); 2[male] 3-11.viii.2007 (MG-35-30); 3[male] 11-27.viii.2007 (MG-35-31); 2[male] 27.viii-6.ix.2007 (MG-35-32); 1[male] 6-13.ix.2007 (MG-35-33); 6[male] 13-20.ix.2007 (MG-35-34). Tulear Prov.: 1[male] Zombitse National Park, near national road, 22[degrees]50.43'S:44[degrees]43.87'E, 26.xi-2. xii.2001, 825 m, R. Harin'Hala, Malaise trap, deciduous spiny forest (MA-02-13B-04); 1[male] Beza Mahafaly Reserve, Parcelle I near research station, 23[degrees]41.19'S:44[degrees]35.46'E, 15.x-10.xi.2001, 165 m, M.E. Irwin, F.D. Parker & R. Harin'Hala, Malaise trap in dry deciduous forest (MA-02-14A-01); 2[male] Mikea Forest, 97 ft, 22[degrees]54.22'S:43[degrees]28.53'E, 6-16.xii.2001, Malaise trap, deciduous dry forest, R. Harin'Hala & M.E. Irwin (MG-18A-05); same except: 2[male] 16-26.xii.2001 (MG-18A-06); 1[male] 26.xi-27.xii.2001 (MG-18A-07); 1[male] 27.xii.2001-6.i.2002 (MG-18A-08); 1[male] 19.ii-1.iii.2002 (MG-18A-14); 1[male] 8-18.iii.2002 (MG-18A-16); 2[male] 18-29.iv.2002 (MG-18A-20); 1[male] 9-19.v.2002 (MG-18A-22); 1[male] 22.vi-2.vii.2002 (MG-18A-26); 1[male] 2-13.vii.2002 (MG-18A-27); 1[male] 23.vii-6.viii.2002 (MG-18A-29); 1[male] 19.viii-3.ix.2002 (MG-18A-31); 3[male] 29.vi-6.vii.2003 (MA-02-18A-60); 2[male] 6-17.vii.2003 (MA-02-18A-61); 2[male] 17-27.vii.2003 (MA-02-18A-62); 5[male] 27.vii-3.viii.2003 (MA-02-18A-63); 2[male] Mikea Forest, 22[degrees]54.80'S:43[degrees]23.93'E, 20-27. xi.2001, 120 ft, R. Harin'Hala & M.E. Irwin, Malaise trap, spiny forest (MG-18B-03); same except: 3[male] 6-16.xii.2001 (MG-18B-05); 1[male] 16-26.xii.2001 (MG-18B-06); 1[male] 6-16.i.2002 (MG-18B-09); 1[male] 16-17.i.2002 (MG-18B-10); 1[male] 28.iii-8.iv.2002 (MG-18B-18); 1[male] 18-29.iv.2002 (MG-18B-20); 1[male] 29.iv-9.v.2002 (MG-18B-21); 1[male] 19-29.v.2002 (MG-18B-23); 1[male] 19.viii-3.ix.2002 (MG-18B-31); 1[male] Beroboka village 45 km NE Morondava, 19[degrees]58.65'S:44[degrees]39.92'E, 26-30.x.2007, 420 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, Antsarongaza gallery forest (MG-45B-06); same except: 1[male] 6-13.xi.2007 (MG-45B-08) (all CAS). Mgjunga Prov.: 1[male] Ampijoroa National Park, 160 km N of MGevatanana on RN 04, 16[degrees]19.16'S:46[degrees]48.80'E, 2-9.vii.2003, 140 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, deciduous forest (MG-25-05); same except: 1[male] 10-17.viii.2003 (MG-25-10) (all CAS); 1[male] 7-14.ix.2003 (MA-25-14) [in spirit], BMSA(DNA)#0042 (BMSA); 1[male] 2-9.xi.2003 (MG-25-22); 1[male] 9-20.xi.2003 (MG25-23); 3[male] 30.xi-8.xii.2003 (MG-25-25); 1[male] 16-27.x.2004 (MG-25-34); 1[male] 21.i-7.ii.2005 (MG-2542); 3[male] 17-19.ii.2005 (MG-25-43). Antananarivo Prov:. 1[male] Ambohitantely, 46 km NE of Ankazobe, 18[degrees]11.88'S:47[degrees]16.89'E, 28.xii.2003-10.i.2004, 2300 ft, R. Harin'Hala & M.E. Irwin, Malaise trap, foret sclerophylle (MG-27-12); 6[male] same except: 10-20.i.2004 (MG-27-13); 1[male] same except: 1-12.ii.2004 (MG-27-15). Majunga Prov.: 2[male] Ambovomamy Belambo, 20 km NW of Port Berger, 15[degrees]27.07'S:47[degrees]36.80'E, 4-14.i.2007, 33 m, R. Harin'Hala, M.E. Irwin & F. Parker, Malaise, secondary growth on white sand (MG-33-01); same except: 3[male] 21-22.i.2007 (MG-33-03); 1[male] 22-27.i.2007 (MG-33-04); 1[male] 7-13. ii.2007 (MG-33-07); 1[male] 13-18.ii.2007 (MG-33-08) (all CAS); 1[male] 18-25.ii.2007 (MG-33-09) [in spirit], BMSA(DNA)#0044 (BMSA); 1[male] 3-10.iii.2007 (MG-33-11); 1[male] 10-18.iii.2007 (MG-33-12); 1[male] 1826.iii.2007 (MG-33-13); 1[male] 6-12.iv.2007 (MG-33-16); 9[male] Besalampy District, Marofototra dry forest, 17 km W of Besalampy, 16[degrees]43.30'S:44[degrees]25.42'E, 24.ix-1.x.2007, 170 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, dry wash in forest (MG-42A-01); 1[male] Besalampy District, Marofototra palm forest 17 km W of Besalampy, 16[degrees]43.30'S:44[degrees]25.42'E, 1-8.x.2007, 35 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, palm trees on sand (MG-42B-02); same except: 6[male] 8-15.x.2007 (MG-42B-03); 13[male] 15-22.x.2007 (MG-42B-04) [1 head missing]; 6$ 22-29.x.2007 (MG-42B-05); 2[male] 29.x-5.xi.2007 (MG-42B-06); 5[male] 2-12.xi.2007 (MG-42B-07); 9[male] 12-19.xi.2007 (MG-42B-08); 9[male] 19-26.xi.2007 (MG-42B-09); 5[male] 26.xi-3.xii.2007 (MG-42B-10); 9[male] 3-10.xii.2007 (MG-42B-11); 8[male] 10-17.xii.2007 (MG-42B-12); 5[male] 17-24.xii.2007, (MG-42B-13); 4[male] Maintirano District, Asondrodava dry forest, 15 km N of Maintirano, 17[degrees]57.92'S:44[degrees]2.13'E, 27.ix-8.x.2007, 200 ft, M.E. Irwin & R. Harin'Hala, Malaise trap in dry forest (MG-43A-01); same except: 4$ 8-15.x.2007 (MG-43A-02); 5[male] 22-29.x.2007 [1 with wing detached and glued to point] (MG-43A-04); 1[male] 29.x-5.xi.2007 (MG-43A-05); 2[male] 5-12.xi.2007 (MG-43A-06); 4[male] 12-17.xi.2007 (MG-43A-07); 1[male] 17.xi-3.xii.2007 (MG-43A-08); 2[male] 3-10.xii.2007 (MG-43A-09); 6[male] 10-17.xii.2007 (MG-43A-10); 26[male] 17-24.xii.2007 (MG-43A-11); 5[male] 24-31.xii.2007 (MG-43A-12); 18[male] 31.xii.2007-7.i.2008 (MG-43A-13); 23[male] 7-14.i.2008 (MG-43A-14); 18[male] 14-21.i.2008 (MG-43A15); 27[male] 21-28.i.2008 (MG-43A-16); 28[male] 28.i-4.ii.2008 (MG-43A-17); 23[male] Maintirano District, Asondrodava dry forest 15 km N of Maintirano, 17[degrees]57.92'S:44[degrees]2.13'E, 27.ix-8.x.2007, 200 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, dry forest at dune (MG-43B-01); same except: 7[male] 8-15.x.2007 (MG-43B-02); 12[male] 15-22.x.2007 (MG-43B-03) [1 head missing]; 11[male] 22-29.x.2007 (MG-43B-04); 5$ 29.x-5.xi.2007 (MG-43B-05); 1[male] 5-12.xi.2007 (MG-43B-06); 3[male] 12-17.xi.2007 (MG-43B-07) (all CAS); 1[male] 10-17. xii.2007 (MG-43B-11) [in spirit], BMSA(DNA)#0053 (BMSA); 3[male] 17-24.xii.2007 (MG-43B-12); 5[male] 24-31.xii.2007 (MG-43B-13); 4[male] 31.xii.2007-7.i.2008 (MG-43B-14) (all CAS); 1[male] 14-21.i.2008 (MG-43B-16) [in spirit], BMSA(DNA)#0055 (BMSA); 2[male] 21-28.i.2008 (MG-43B-17). Toamasina Prov.: 1[male] Mobot site, Analalava, 7 km SW of Foulpointe, 17[degrees]42'32"S:49[degrees]27'29"E, 15-21.ix.2007, 80 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, low altitude dense humid forest (MG-37A-01) (CAS).

Distribution (Fig. 98): The most widely distributed species of the genus in Madagascar, distributed in six vegetation types (Appendix II), in the Dry Deciduous Forest, Central Highlands and Evergreen Rainforest biomes. Occurring in nine biogeographical zones and all bioclimatic zones (Figs 105-107; Tables 1-3).

Curtonotum boeny Tsacas, 1974

Figs 10, 23, 36, 46, 49, 52, 88, 101

Curtonotum boeny: Tsacas 1974: 712; figs 5a-9 (p. 713), fig. 8c (p. 718). Type locality: "Madagascar: Ambato-Boeny".

Differential diagnosis: This species is closely related to C. griveaudi sp. n., differing in the colour of the frons and antennae, the shape of the wing and the shape of the male terminalia. Both share the peculiar subtriangular form of the epandrium with the lateral setation reduced to two long and strong setae originating from the blunt point formed by the ventral margin. Curtonotum boeny differs from C. griveaudi sp. n., however, in having the hypandrial arm narrowed apically, the surstylus shorter and wider basally, the presence and position of the conspicuous internal bulge on the phallus and the apical shape of the basiphallus, with the more pronounced finger-like medial projection. The ranges of the two species overlap, but it is not known whether they occur sympatrically.

Redescription:

Male (primarily based on field-pinned HT).

As redescribed for C. balachowskyi, differing in the following respects:

Measurements: Overall length 3.2-4.5 mm (n = 6, N-T); length of head and thorax combined 2.1 mm; length of thorax and scutellum combined 1.95 mm; wing length 3.2 mm (HT).

Head (Figs 10, 23). Eye prominent, eye height/length ratio: 9:5 (HT); frons (Fig. 23), slightly wider than long, frons length/width ratio: 50:55 (HT), markedly wider at vertex than at ventral margin; arista with 8 long dorsal branches and 3 ventral branches in addition to terminal fork; face with narrow silver fascia (adjacent to eye margin); 1 pair of weak vibrissae inserted on posterior lateral margin and 10 much finer setae bordering genal groove; occiput grey pruinose throughout; gena narrow, eye height/ genal height ratio: 9:1 (HT), silver pruinose, slightly dirty yellow beyond basal angle; palpus brown.

Thorax (Fig. 10). Mesonotum with two narrow, weakly-defined, parallel, medial chestnut-brown pruinose vittae on dorsal surface, extending from anterior margin to region of anterior dorsocentral seta socket; anterior dorsocentral seta much shorter and finer than medial scutellar seta; reclinate supra-alar seta, slightly shorter than posterior dorsocentral seta; thoracic pleurae silver-grey, yellow pruinose in their centres; postpronotum silver-grey pruinose, with 8-10 fine black-brown setulae; anepisternum silver-grey pruinose, with yellow pruinose patches medially, with 19-20 fine setulae, the 5 larger grouped together at posterodorsal margin; katepisternum with dorsal katepisternal seta ca 0.3 length of ventral katepisternal seta, with 18 short, fine setulae at base and along posterior margin.

Scutellum. As in mesonotum; two pairs of strong scutellar setae, 1 weak basal scutellar setula and 1 weak intermediate scutellar setula, the latter inserted equidistant between lateral and medial scutellar setae.

Legs. Fore coxa with 8-10 brown setulae on anterior surface; hind coxa with lateral black seta; fore tibia with ctenidium of 8 short spinules.

Wing (as in Fig. 36). Veins brown, membrane hyaline with very faint brown infuscation, slightly darker in region of dm-cu crossvein; dm-cu crossvein obliquely angled.

Abdomen. Tergite 1 simple, devoid of maculae; tergite 2 with oblique, subovoid brown-black pruinose dorsolateral macula on either side only; tergites 3-5 with large broad, V-shaped concolourous median fascia and large concolourous T-shaped dorsolateral maculae, these close to, but not fully merging with, median fascia; lateral margin of tergites 2-5 with subelliptical concolourous macula in basal half; sternite 4 quadrate, weakly sclerotised; sternite 5 slightly longer than sternite 4, lateral margins rounded, with oblong brown macula on either side, both unmodified, with sparse brown setulae arranged in irregular rows, those along apical margins of sternite 5 slightly longer and stronger; sternite 6 (Fig. 88) subquadrate (may appear narrower than Fig. 88 in undissected specimens), slightly expanded medially, with moderately deep, subtriangular excision apically, brown in apical 0.8, clothed in closely-packed, long, curved, overlapping brown setulae in apical 0.8.

Terminalia (Figs 46, 49, 52). Hypandrium (Fig. 46, hy) rather flattened and long, with broad-based, rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (subtriangular in profile); hypandrial arms of even width (viewed laterally), with 2 setulae proximal to postgonite, the more lateral ventrally directed, the medial ventromedially directed (obscured by epandrium on Fig. 46), sclerotised area of medial lobes (viewed dorsally), with slightly rounded margins, not overlapping (not convex); postgonite (Fig. 46, pg) long, thin and straight; epandrium (Fig. 46, ep) subtriangular (in lateral view), with deep, oblique excavation along ventral margin, ventral lobe forming a blunt point from which two very strong, long setae originate; cercus (Fig. 46, ce) not prominent, longest setae as long as medium setae on dorsal margin of epandrium; surstylus (Fig. 46, ss) long, widest basally, slightly curved in apical 2/3; phallus (as in Figs 49,ph, bp, dp; 52, bp, dp) C-shaped, moderately sclerotised, brown; phallapodeme (Fig. 49, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, broad, subtriangular projections in basal 0.4, bi-furcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 49, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 49, bp) broader in basal half, slightly narrowed in apical half, with large, conspicuous internal bulge at point of first bend in basal fifth; apical section (Fig. 52, bp) moderately broad, right lateral margin with raised keel, projecting medially as long, finger-like projection; distiphallus (Figs 49, 52, dp) long, with slight curve at midlength, very narrow (viewed laterally and dorsally).

[FIGURES 9-13 OMITTED]

Variation: The frons appears slightly darker in ex-alcohol specimens.

Holotype (examined): [male] MADAGASCAR: "MADAGASCAR Maj. / Ambato-Boeni / 23.VI.[19]58 F. KEISER [pink paper] // HOLOTYPE [red card] // CURTONOTUM / boeny n.sp. / Holotype [male] / L. TSACAS DET. 1973 [printed & handwritten] // MUSEUM PARIS // Curtonotum / boeny [male] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit 2008" (MNHN). In excellent condition; pinned and double-mounted on card triangle; right wing detached and glued to point; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "Curtonotum / boeny [female] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit 2008-2010"): MADAGASCAR: 1[female] same labels as holotype, except: "// ALLOTYPE [red card] // CURTONOTUM / boeny n.sp. / Allotype [female] / L. TSACAS DET. 1973 [printed & handwritten]" [left wing detached and glued to card] (MNHN); 2[female] same except: "// PARATYPE [red card] // CURTONOTUM / boeny n.sp. / Paratype [female] / L. TSACAS DET. 1973 [printed & handwritten]" (NHMB).

Notes: Tsacas (1974: 713) cited the date of collection of the type series as "23.v.58", but the labels actually read "23.VI.58". He further noted that the holotype and 1 paratype were deposited in MNHN and that the allotype and 1 paratype were deposited in NHMB. In truth, the holotype and allotype are deposited in MNHN and 2 paratypes in NHMB.

Other material examined (all labelled: "Curtonotum / boeny [male] [or [female]] / Tsacas, 1974 / det. A.H. Kirk-Spriggs 2010"): MADAGASCAR: Mahajanga Prov.: 4[male] Parc National Tsingy de Memaraha, 3.4 km 93[degrees] E Bekopaka, Tombeau Vazimba, 19[degrees]08'31"S:44[degrees]49'41"E, 50 m, 6-10.xi.2001, leg. [B.] Fisher, [C.] Griswold et al., tropical dry forest, Malaise trap (MAD49), BLF4233 (CAS).

Distribution (Fig. 101): Occurring in the Wooded Grassland-Bushland and Western Dry Forest vegetation types, in the Dry Deciduous Forest biome. In the North West and West biogeographical zones and Dry bioclimatic zone (Figs 105-107; Tables 1-3; Appendix II).

Curtonotum coronaeformis sp. n.

Figs 6, 19, 32, 66, 69, 72, 83, 97

Etymology: From Latin corona (crown) and formis (in the form of), and refers to the crown-like lateral extension of the distiphallus of this species.

Differential diagnosis: This species is closely related to C. parkeri sp. n.; the shape, maculae and setation of abdominal sternites 4 and 5 are virtually identical, the apical region of the basiphallus is markedly expanded in both species, with very similar left and right raised keels and both share the raised and spinose right sclerotised area of the basiphallus. In C. coronaeformis sp. n., however, the basiphallus is less markedly narrowed medially and less expanded in the apical third, and the raised and spinose right sclerotised area of the distiphallus is conspicuously developed, with a series of regular to irregular spines. The two species occur sympatrically.

Description:

Male (primarily based on ex spirit-preserved HT).

As redescribed for C. balachowskyi, differing in the following respects:

Measurements: Overall length unknown; length of head and thorax combined 2.9 mm; length of thorax and scutellum combined 2.4 mm; wing length 3.7 mm.

Head (Figs 6, 19). Compound eye prominent, probably green-brown iridescent in living examples, eye height/length ratio: 10:7 (HT); frons (Fig. 19), slightly wider than long, frons length/width ratio: 5 :6 (HT), markedly wider at vertex than at ventral margin, ground colour pale yellow to pale brown, darker brown towards vertex and between orbital plates forming conspicuous vittae that reach ventral margin; orbital plates and ocellar triangle golden-grey pruinose; orbital plates extending from vertex of head to ca 0.8 length of frons, margins regular; antennal scape and pedicel dirty pale brown, silver-grey pruinose, flagellomere 1 very long, ca 2.5x longer than wide, apex evenly-rounded, yellow basally, darkened apically, yellow-grey pruinose, arista with 8 or 9 long dorsal branches and 3 ventral branches in addition to terminal fork; lunule and face silver-grey pruinose, yellow pruinose beneath flagellomere 1; gena narrow, eye height/genal height ratio: 10:1 (HT), silver pruinose throughout; palpus pale brown.

Thorax (Fig. 6). Mesonotum as described for C. boeny; acrostichal setae, much shorter than anterior dorsocentral seta; supra-alar seta, slightly shorter than posterior dorsocentral seta; postalar setae longer and stronger than acrostichal setae; postpronotum yellow-grey pruinose, with 6 finer black-brown setulae; anepisternum silver-grey pruinose, with yellow pruinose patches medially, surface with 18 fine setulae, some larger and arranged in 2 groups of 3 and 4; katepisternum silver-grey to silver-yellow pruinose, with 13 short, fine setulae at base and along posterior margin.

Scutellum. As in mesonotum, with very faint medial brown pruinose vitta basally (under some lights), slightly paler yellow pruinose at posterior margin; weak intermediate scutellar setula inserted at 0.8 distance between medial and lateral scutellar setulae.

Legs. Fore coxa with 22 brown setulae on anterior surface; fore tibia with ctenidium of 10 or 11 short, sharp black spinules.

Wing (as in Fig. 32). Veins chestnut-brown, membrane very faintly infuscate brown throughout, very slightly darker in r} and in region of dm-cu crossvein; dm-cu crossvein oblique, in shape of uninterrupted arc.

Abdomen. Tergites 3-5 with broad V-shaped, median fascia adjoining and slightly merging with large, concolourous T-shaped dorsolateral maculae; lateral margin of tergites 2-5 with subelliptical concolourous macula in basal half to %; sternite 4 with posterior and lateral margins evenly rounded, apical margin straight; sternite 5 rectangular, similarly shaped to sternite 4, slightly longer and wider medially, with 2 small ovoid basomedial maculae, sternites 4 and 5 unmodified, with long, dense, brown setulae arranged in irregular rows, those along lateral margins longer and stronger; sternite 6 (Fig. 83) narrowed basally, evenly rounded laterally (may appear narrower than Fig. 83 in undissected specimens), with broad, deep apical excision and brown maculae medially and fascia laterally, merging apically, clothed in long, black, irregular, medially-directed, brown setulae in apical 0.8, those at apical margin longer and more prominent.

Terminalia (Figs 66, 69, 72). Hypandrium (Fig. 66, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (rounded to slightly angulate in profile); hypandrial arms constricted in apical 2/ (viewed laterally), with 2 setulae proximal to postgonite, the more lateral ventrally directed, the medial ventromedially directed (obscured by epandrium on Fig. 66), sclerotised area of medial lobes (viewed dorsally), with margins evenly rounded, convex, closely abutting, not overlapping; postgonite (Fig. 66, pg) long, thin, spindle-like; epandrium (Fig. 66, ep) broad (viewed laterally), evenly-rounded on dorsal margin, posterior margin slightly angled, ventral margin with extensive row of long, regular to irregular, apically-directed setae; cercus (Fig. 66, ce) not prominent, longest setae longer than setae on dorsal margin of epandrium; surstylus (Fig. 66, ss) long and narrow, slightly curved in apical 2; phallus (as in Figs 69, ph, bp, dp, 72, bp, dp) C-shaped, moderately sclerotised, brown; phallapodeme (Fig. 69, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, narrow, rounded projections in basal 0.4, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 69, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 69, bp) broad basally, narrowed just beyond midlength; apical section (Fig. 72, bp) expanded and broad, right lateral margin with ventral projection, left margin with distinct raised keel; distiphallus (Figs 69, 72, dp) long, narrow and curved, with extensive broad membranous section, basolaterally with upturned, sclerotised, crown-like projection (subtriangular viewed laterally).

[FIGURES 14-26 OMITTED]

Variation: The size and number of teeth and smaller serrations at the margin of the upturned, sclerotised, crown-like projection of the distiphallus are variable. This is here interpreted as intraspecific variation only, since other terminalia characters are constant.

Holotype [male] "MADAGASCAR: Province / Fianarantsoa, near Isalo / National Park, in dry wash / east of Interpretive Center / 22[degrees]37.60'S, 45[degrees]21.49'E / 23.ii-5.iii.2002, 885 m / M.E. Irwin & R. Harin'Hala / Malaise trap in open area / MA-02-11B-57 // HOLOTYPE [male] / Curtonotum / coronaeformis sp. n. / A.H. Kirk-Spriggs 2010 [red card]" (CAS). In good condition; card-pointed; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "PARATYPE $ / Curtonotum / coronaeformis sp. n. / A.H. Kirk-Spriggs 2010 [blue card]"): 3[male] same data as holotype, except (1 labelled: "BMSA(DNA)#0049") [1 right wing detached and glued to card]; same except: 1[male] "12-22.vi.2002, MA-02-11B-32"; 1[male] "6-14.xii.2002, MA-02-11B-49" (all CAS).

Distribution (Fig. 97): Apparently confined to the Wooded Grassland-Bushland vegetation type, in the Central Highlands biome. In the Central biogeographical zone and Subarid bioclimatic zone (Figs 105-107; Tables 1-3; Appendix II).

Curtonotum gladiiformis sp. n.

Figs 8, 21, 34, 41, 43, 45, 85, 99

Etymology: From Latin gladius (sword) and formis (in the form of), and refers to the sword-like shape of the distiphallus of this species.

Differential diagnosis: This species is probably most closely related to C. balachowskyi Tsacas. Both share the similarly-shaped scimitar-like distiphallus, with the incurved basal area (viewed laterally) and the subquadrate sternite 6, with a very shallow apical excision. Curtonotum gladiiformis differs from C. balachowskyi, however, in having the lateral margin of the apical border of the basiphallus developed into a medial and submedial spine, as opposed to C. balachowskyi, in which the right lateral border forms a distinct, angulate fold, and in having the setae on sternite 6 much longer on the apical margin.

Description:

Male (primarily based on ex spirit-preserved HT).

As redescribed for C. balachowskyi, differing in the following respects:

Measurements: Overall length unknown; length of head and thorax combined 2.9 mm; length of thorax and scutellum combined 2.6 mm (HT); wing 4.4 mm long (n = 1, PT).

Head (Figs 8, 21). Eye prominent, eye height/length ratio: 11:7 (HT); frons (Fig. 21), slightly wider than long, frons length/width ratio: 7:8 (HT), ground colour pale brown, darker brown towards vertex and between orbital plates forming conspicuous vittae that reach ventral margin; orbital plates and ocellar triangle golden-grey pruinose; ocelli clear brown; orbital plates extending from vertex of head to 0.8 length of frons, lateral margins with narrow silver pruinose fascia (adjacent to eye margin), slightly wider at antennal insertions; posterior orbital seta moderately strong, slightly longer than outer vertical seta; antennal scape and pedicel brown-grey pruinose, flagellomere 1 yellow pruinose basally, pitchy brown, grey pruinose apically, arista with 8 or 9 long dorsal branches and 3 or 4 ventral branches in addition to terminal fork; lunula yellow-grey pruinose; face with narrow silver fascia (adjacent to eye margin); occiput grey pruinose; gena narrow, eye height/genal height ratio: 10:1 (HT), silver pruinose throughout, very slightly darker beyond basal angle; palpus pale brown, brown microtrichose.

Thorax (Fig. 8). Mesonotum as described for C. boeny; presutural seta (missing on holotype and paratypes); anterior notopleural seta markedly longer than posterior; supraalar seta shorter than posterior dorsocentral seta; postalar setae longer and stronger than acrostichal setae; postpronotum silver-yellow pruinose, with 11 finer black-brown setulae; anepisternum silver-grey pruinose, silver-yellow pruinose medially, surface with 24 fine setulae, some larger and arranged in 2 groups of 3; anepimeron, laterotergite and meron silver-grey to yellow-grey pruinose; katepisternum silver-grey to yellow-grey pruinose with darker grey macula in anterior half, dorsal katepisternal setae ca 0.4 length of ventral katepisternal setae, with 11 short, fine setulae at base and along posterior margin.

Scutellum. As in mesonotum, slightly paler yellow pruinose at posterior margin; weaker intermediate scutellar setula inserted at 0.8 distance between medial and lateral scutellar setulae.

Legs. Fore coxa with 9 brown setulae on anterior surface; mid coxa with 8 brown setulae; fore tibia with ctenidium of 12 spinules.

Wing (as in Fig. 34). Broad basally, tip evenly rounded, veins chestnut-brown, membrane very faintly infuscate brown throughout, very slightly darker in region of dm-cu crossvein; dm-cu crossvein evenly arched; haltere dirty white.

Abdomen. Tergite 1 simple, devoid of maculae; tergite 2 with oblique, subrectangular brown-black pruinose dorsolateral macula on either side and faint medial fascia; tergites 3-5 with broad, V-shaped concolourous median fasciae and well separated and extensive concolourous T-shaped dorsolateral macula, these close, but not merging with medial fascia, lateral margin of tergites 2-5 with subquadrate concolourous macula in basal 2/3; sternite 4, weakly sclerotised, quadrate, sides evenly narrowing to straight apical margin; sternite 5 similar, but wider and longer, both unmodified, with long, brown, sparse setulae arranged in irregular rows, lateral setulae longer and stronger; sternite 6 (Fig. 85) much broader than long (may appear narrower than Fig. 85 in undissected specimens), with very shallow apical excision and faint, transverse, brown maculae apically, clothed in extremely long, sparse, black, irregular setulae, especially along apical margin.

[FIGURES 27-39 OMITTED]

[FIGURES 40-45 OMITTED]

Terminalia (Figs 41, 43, 45). Hypandrium (Fig. 41, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (rounded to slightly angulate in profile); hypandrial arm constricted medially (viewed laterally), with 2 setulae proximal to postgonite, the more lateral strong, ventrally-directed, the medial much smaller and weaker (or absent) (obscured by epandrium on Fig. 41), sclerotised area of medial lobes (viewed dorsally), with margins evenly rounded, convex medially, closely abutting, not overlapping; postgonite (Fig. 41, pg) long, thin and straight, with slight undulating anterior margin; epandrium (Fig. 41, ep) broader dorsally than ventrally (viewed laterally), evenly rounded on dorsal margin, posterior margin slightly angled, ventral margin with extensive row of long, regular to irregular, ventrally-directed setae; cercus (Fig. 41, ce) not prominent, longest setae longer than dorsal setae of epandrium; surstylus (Fig. 41, ss) long, widest basally, slightly curved in apical 22; phallus (as in Figs 43, ph, bp, dp; 45, bp, dp) C-shaped, very large, moderately sclerotised, brown; phallapodeme (Fig. 43, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, narrow, subtriangular projections in basal 0.4, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 43, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 43, bp) broad in basal region, with conspicuous internal bulge, then narrowed, moderately constricted in ca apical 2/3 (viewed dorsally); apical section (Fig. 45, bp) broad, right lateral margin evenly rounded and flat, developed into a bluntly-pointed ventromedial spine, with much smaller spine basolaterally; distiphallus (Figs 43, 45, dp) moderately sclerotised, extremely long, sword-like, with conspicuous curved indentation in basal third (viewed laterally), sclerotised area subdivided basally, with triangular window.

Variation: In the two PT the basiphallus is not as markedly constricted in the apical 2/3 as in Fig. 43 and this may be the result of slight distortion in phallus of the HT.

Holotype: [male] "MADAGASCAR: Province / Fianarantsoa, near Isalo / National Park, in dry wash / east of Interpretive Center / 22[degrees]37.60'S, 45[degrees]21.49'E / 28.iii-9.iv.2003, 885 m / M.E. Irwin & R. Harin'Hala / Malaise trap in open area / MA-02-11B-60 // HOLOTYPE [male] / Curtonotum / gladiiformis sp. n. / A.H. Kirk-Spriggs 2010 [red card]" (CAS). In good condition, some head setae missing; card-pointed; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "PARATYPE [male] / Curtonotum / gladiiformis sp. n. / A.H. Kirk-Spriggs 2010 [blue card]"): 1[male] same data as holotype, except: "3-10.ii.2002, MA-02-11B-14" [right wing detached and glued to card]; 1[male] "6-14.xii.2002, MA-02-11B-49 // BMSA(DNA)#0050" (both CAS).

Distribution (Fig. 99): Apparently confined to the Wooded Grassland-Bushland vegetation type, in the Central Highlands biome. In the Central biogeographical zone and Subarid bioclimatic zone (Figs 105-107; Tables 1-3; Appendix II).

Curtonotum griveaudi sp. n.

Figs 11, 24, 37, 47, 50, 53, 89, 102

Etymology: The species is named in honour of Paul Elexis Jacques Griveaud (1907-1980), in recognition of his contribution to Madagascan entomology.

Differential diagnosis: This species is closely related to C. boeny, differing in the colour of the frons and antennae, the shape of the wing and the shape of the male terminalia. Both share the peculiar subtriangular form of the epandrium, with the lateral setation reduced to two long, strong setae originating from the blunt point formed by the ventral margin. Curtonotum griveaudi sp. n. differs from C. boeny, however, in having the hypandrial arm much broader apically and the surstylus considerably longer and narrower basally. The species further differ in the presence and position of the low internal bulge of the phallus, with the basal section of the phallus quite straight, and the apical shape of the basiphallus, with its reduced lateral keel and short, spine-like ventromedial projection. The ranges of the two species overlap, but it is not known whether they occur sympatrically.

Description:

Male (primarily based on ex spirit-preserved HT).

As redescribed for C. balachowskyi, differing in the following respects:

Measurements: Overall length unknown; length of head and thorax combined 2.2 mm; length of thorax and scutellum combined 2.2 mm (HT); wing length 3.5 mm (n = 1, N-T).

Head (Figs 11, 24). Eye prominent, eye height/length ratio: 10:6 (HT); frons (Fig. 24), markedly wider at vertex than at ventral margin, frons length/width ratio: 1:5 (HT), ground colour pale brown to dark brown, darker brown towards vertex and between orbital plates forming conspicuous vittae that reach ventral margin; ocellar triangle golden-grey pruinose, ocelli clear brown; orbital plates extending from vertex of head to 0.8 length of frons, markedly wider at socket of posterior orbital seta; lateral margins with narrow silver pruinose fascia (adjacent to eye margin), slightly wider at antennal insertions; posterior orbital seta moderately strong, slightly longer than outer vertical seta; antennal scape and pedicel brown-grey pruinose, flagellomere 1 yellow pruinose basally, pitchy brown, grey pruinose apically, arista with 10 long dorsal branches and 4 ventral branches in addition to terminal fork; lunule brown, face with narrow silver fascia (adjacent to eye margin); occiput grey pruinose; gena narrow, eye height/genal height ratio: 10:1 (HT), silver pruinose throughout, very slightly darker beyond basal angle; palpus pale brown, brown microtrichose.

Thorax (Fig. 11). Mesonotum as described for C. boeny; anterior dorsocentral setae much shorter and finer than medial scutellar seta; postalar setae moderately strong, slightly longer and stronger than acrostichal setae; postpronotum silver-grey pruinose, with 7 or 8 finer black-brown setulae; anepisternum silver-grey pruinose, with yellow pruinose patches medially, surface with 28 fine setulae, some larger and arranged in 2 groups of 3 and 4; anepimeron, laterotergite and meron silver-grey to yellow pruinose; katepisternum silver-grey to yellow pruinose, with dorsal katepisternal setae ca 0.2 length of ventral katepisternal setae, with 18 short, fine setulae at base and along posterior margin.

Scutellum. As in mesonotum, with very faint medial brown pruinose vitta basally (under some lights), slightly paler yellow pruinose at posterior margin; weak intermediate scutellar setula inserted closer to lateral than medial scutellar setae.

Legs. Fore coxa with 18 brown setulae on anterior surface; mid coxa with 16 brown setulae; fore tibia with ctenidium of 10 spinules.

Wing (as in Fig. 37). Veins brown, membrane hyaline with very faint brown infuscation, very slightly darker in posterior half of r1 and anterior half of r2+3 and in region of dm-cu crossvein; dm-cu crossvein strongly, obliquely angled in even arc.

Abdomen. Tergite 1 simple, devoid of maculae; tergite 2 with oblique, subovoid brown-black pruinose dorsolateral macula on either side only; tergites 3-5 with large, broad V-shaped median fascia and large concolourous T-shaped dorsolateral macula, these close or adjoining, but not fully merging with median fascia; lateral margin of tergites 2-5 with subelliptical concolourous macula in basal half; sternite 4, long, rectangular, weakly sclerotised; sternite 5 same length as sternite 4, lateral margins curved, with oblong brown macula on either side, both unmodified, with very sparse brown setulae arranged in irregular rows, those along lateral margins of sternite 5 slightly longer and stronger; sternite 6 (Fig. 89) rather broad apically, narrow basally (may appear narrower than Fig. 89 in undissected specimens), with moderately shallow, subtriangular excision apically, brown in apical 0.8, clothed in sparse, long, straight, overlapping brown setulae in apical 0.8.

Terminalia (Figs 47, 50, 53). Hypandrium (Fig. 47, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (subtriangular in profile); hypandrial arms expanded apically, with sclerotised ventrally-directed arc (viewed laterally), setulae absent, sclerotised area of medial lobes (viewed dorsally), with slightly rounded margins, not overlapping (not convex); postgonite (Fig. 47, pg) long, thin and straight; epandrium (Fig. 47, ep) subtriangular (in lateral view), with deep, oblique semicircular excavation along ventral margin, ventral lobe forming a blunt point, from which two very strong, long setae originate; cercus (Fig. 47, ce) not prominent, longest setae as long as medium setae on dorsal margin of epandrium; surstylus (Fig. 47, ss) drawn out, very long and narrow, slightly wider basally and curved in apical 0.8; phallus (as in Figs 50, ph, bp, dp, 53, bp, dp) C-shaped, weakly sclerotised, pale brown; phallapodeme (Fig. 50, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, broad, subtriangular projections in basal 0.4, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 50, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 50, bp) broad and straight in basal half, broadest at first bend, with slight internal bulge, then markedly narrowed (narrowest at second bend); apical section (Fig. 53, bp) very broad, right margin laterally expanded and evenly rounded, with slightly raised margin, left lateral margin with distinct raised fold, projecting medially as short, spine-like ventromedial projection; distiphallus (Figs 50, 53, dp) long, evenly curved, scimitar-like (viewed laterally), narrow (viewed dorsally).

[FIGURES 46-54 OMITTED]

Variation: The frons is darker in some specimens, almost chestnut-brown; even in specimens with a paler brown frons the two longitudinal brown vittae are strikingly apparent.

Holotype: [male] "Madagascar: Majunga / Prov., Maintirano District / Asondrodava dry forest / 15km N of Maintirano / 17[degrees]57.92'S 44[degrees]2.13'E / 24-31.xii.2007, 200 ft / M.E. Irwin & R. Harin'Hala / Malaise trap, dry forest / at dune, MG-43B-13 // HOLOTYPE [male] / Curtonotum / griveaudi sp. n. / A.H. Kirk-Spriggs 2011 [red card]" (CAS). In good condition, right foreleg missing; card-pointed; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "PARATYPE [male] [or [female]] / Curtonotum / griveaudi sp. n. / A.H. Kirk-Spriggs 2011 [blue card]"): 1[male] "Madagascar: Tulear / Province, Mikea Forest, / NW of Manombo / 22[degrees]54.22'S, 43[degrees]28.53'E / 6-17.vii.2003, 30 m / M.E. Irwin & R. Harin'Hala / Malaise trap in deciduous / dry forest / MA-02-18A-61"; 1[male] "Madagascar: Province / Fianarantsoa, near Isalo / National Park, in dry wash / east of Interpretive Center / 22[degrees]37.60'S:45[degrees]21.49'E / 6-14.xii.2002, 885 m / M.E. Irwin & R. Harin'Hala / Malaise trap in open area / MA-02-11B-49"; 1[male] "Madagascar: Tulear / Province, Beza Mahafaly / Reserve, Parcelle I near / research station / 23[degrees]41.19'S:44[degrees]35.46'E / 9-20.ix.2002, 165 m / R. Harin'Hala, Malaise / trap in dry deciduous forest / MA-02-14A-35 // BMSA(DNA)#0047" (all CAS); 1[male] 2[female] "Madagascar NE, Sambava Beach, 14.iv.1991, A. Freidberg & Fini Kaplan" [$ with phallus missing from vial; left wing detached and glued to card] (all TAU).

Other material examined (labelled: "Curtonotum / griveaudi [male] / sp. n. / A.H. Kirk-Spriggs 2011"): MADAGASCAR: Majunga Prov.: 1[male] Maintirano District, Asondrodava dry forest 15 km N of Maintirano, 17[degrees]57.92'S:44[degrees]2.13'E, 10-17.xii.2007, 200 ft, M.E. Irwin & R. Harin'Hala, Malaise trap, dry forest at dune (MG-43B-11) [in spirit], BMSA(DNA)#0054 (BMSA).

Distribution (Fig. 102): Occurs in the Wooded Grassland-Bushland and South Western Dry Spiny Forest-Thicket vegetation types, in the Arid Spiny Bush, Central Highlands and Dry Deciduous Forest biomes. In the West and South biogeographical zones and Dry and Subarid bioclimatic zones (Figs 105-107; Tables 1-3; Appendix II).

Curtonotum irwini sp. n.

Figs 4, 17, 30, 64, 67, 70, 81, 95

Etymology: The species is named in honour of Michael Edward Irwin, in recognition of his contribution to our knowledge of Madagascan Diptera.

Differential diagnosis: This species is probably most closely related to C.parkeri sp. n. and C. coronaeformis sp. n. All share the extensively-developed sclerotised area of the distiphallus and sternite 6 is very similar in all three species. It differs from C. parkeri and C. coronaeformis, however, in lacking a raised lateral, spinose ridge on the distiphallus and in having the right lateral margin of the basiphallus developed into a long, curved, ventrally-directed spine, with smaller spine medially.

Description:

Male (primarily based on ex spirit-preserved HT).

As redescribed for C. balachowskyi, differing in the following respects:

Measurements: Overall length 3.85-4.55 mm (n = 8, PT); length of head and thorax combined 2.5 mm; length of thorax and scutellum combined 2.3 mm (HT); wing length 3.2 mm (n = 1, PT).

Head (Figs 4, 17). As described for C. gladiiformis sp. n., except: eye height/length ratio: 12:8 (HT); frons (Fig. 17), frons length/width ratio: 6:7 (HT); posterior orbital seta moderately strong, slightly shorter than outer vertical seta; arista with 9 or 10 long dorsal branches and 4-6 ventral branches in addition to terminal fork; gena narrow, eye height/genal height ratio: 12:1 (HT), silver pruinose, slightly darker beyond basal angle; palpus pale brown.

Thorax (Fig. 4). Mesonotum as described for C. boeny, except: presutural seta moderately strong, shorter and weaker than posterior notopleural seta; postpronotum with 7 finer black-brown setulae; anepisternum silver-grey pruinose, with silver-yellow pruinose areas, surface with 18 fine setulae, some larger and arranged in 2 groups of 3 and 2; anepimeron, laterotergite and meron silver-grey to silver-yellow pruinose; katepisternum silver-grey to silver-yellow pruinose, with darker macula in anterior half, with dorsal katepisternal setae ca 0.3 length of ventral katepisternal setae, with 18 short, fine setulae at base and along posterior margin.

Scutellum. As described for C. gladiiformis sp. n.

Legs. Fore coxa with 13 brown setulae on anterior surface; mid coxa with 7 brown setulae; fore tibia with ctenidium of 14 spinules.

Wing (as in Fig. 30). Veins chestnut-brown, membrane very faintly infuscate brown throughout, very slightly darker in region of dm-cu crossvein; dm-cu crossvein acutely angled with interrupted arc; haltere dirty white.

Abdomen. Tergite 1 simple, devoid of maculae; tergite 2 with oblique, subovoid brownblack pruinose dorsolateral macula on either side only; tergites 3-5 with large broad, V-shaped concolourous median fascia and large concolourous T-shaped dorsolateral macula, these close or adjoining, but not fully merging with median fascia; lateral margin of tergites 2-5 with subelliptical concolourous macula in basal half; sternites 4-5, as described for C. coronaeformis sp. n.; sternite 6 (Fig. 81), narrowed basally, evenly rounded laterally (may appear narrower than Fig. 81 in undissected specimens), with broad, shallow apical excision and brown maculae medially and fascia laterally, merging apically, clothed in long, black, irregular, apically-directed, brown setulae in apical half, those at apical margin much longer and more prominent.

Terminalia (Figs 64, 67, 70). Hypandrium (Fig. 64, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (rounded to slightly angulate in profile); hypandrial arm constricted apically (viewed laterally), with 2 setulae proximal to postgonite, the more lateral strong, ventrally directed, the medial much smaller and weaker (obscured by epandrium on Fig. 64), sclerotised area of medial lobes (viewed dorsally), with margins evenly rounded, convex medially, closely abutting, not overlapping; postgonite (obscured by surstylus on Fig. 64) long, thin and straight, with slight undulating anterior margin; epandrium (Fig. 64, ep) broader dorsally than ventrally (viewed laterally), evenly rounded on dorsal margin, posterior margin slightly angled, ventral margin with extensive row of long, regular to irregular, apically-directed setae; cercus (Fig. 64, ce) not prominent, longest setae as long as setae on dorsal margin of epandrium; surstylus (Fig. 64, ss) long, widest basally, slightly curved in apical 2/3; phallus (as in Figs 67, ph, bp, dp, 70, bp, dp) C-shaped, heavily sclerotised, brown; phallapodeme (Fig. 67, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, narrow, subtriangular projections in basal 0.4, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 67, ea) free, duct inserted at junction of phallapodeme and basiphallus; basiphallus (Fig. 67, bp) broad in ca basal half, with slight internal bulge at midlength, then markedly narrowed, strongly constricted in ca apical 2 (viewed dorsally); apical section (Fig. 70, bp) moderately broad, right lateral margin evenly rounded, left lateral margin developed into long, curved ventrallydirected spine, with smaller spine medially; distiphallus (Figs 67, 70, dp) broad basally, sclerotised area extensive, abruptly narrowed towards apex, basal section with membranous window, apical section narrow and spindle-like.

Variation: The length of the larger, curved ventrally-directed spine of the distiphallus is variable in some specimens. Other terminalia characters are constant and this is here interpreted as intraspecific variation only.

Holotype: [male] "MADAGASCAR: Tulear Province / Zombitse National Park / near ANGAP office, 840 m / 22[degrees]53.19'S:44[degrees]41.53'E / 28.ii-6.iii.2002 / California Acad of Sciences / R. Harin'Hala, Malaise trap /deciduous spiny forest / MA-02-13A-18 // HOLOTYPE [male] / Curtonotum / irwini sp. n. / A.H. Kirk-Spriggs 2010 [red card]" (CAS). In fair condition, wings slightly tattered and mesonotum rubbed; card-pointed; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratypes (all labelled: "PARATYPE [male] / Curtonotum / irwini sp. n. / A.H. Kirk-Spriggs 2010 [blue card]"): MADAGASCAR: same data as holotype, except: 2[male] "15.x-9.xi.2001, MA-02-13A-01"; 12[male] "9-19.xi.2001, MA-02-13A-02"; 3[male] "19-26.xi.2001, MA-02-13A-03"; 3[male] "26.xi-2.xii.2001, MA-02-13A-04"; 6[male] "2-7.xii.2001, MA-02-13A-05"; 1[male] "7-14.xii.2001, MA-02-13A-06"; 5[male] "14-16. xii.2001, MA-02-13A-07"; 2[male] "16-22.xii.2001, MA-02-13A-08"; 1[male] "22-29.xii.2001, MA-02-13A09"; 8[male] "29.xii.2001-5.i.2002, MA-02-13A-10"; 4[male] "5-12.i.2002, MA-02-13A-11"; 1[male] "19-26.i.2002, MA-02-13A-13"; 5[male] "5-14.ii.2002, MA-02-13A-15"; 12[male] "14-21.ii.2002, MA-02-13A-16"; 7[male] same data as holotype; 1[male] "6-13.iii.2002, MA-02-13A-19"; 1[male] "13-20.iii.2002, MA-02-13A-20"; 9[male] "27. iii-3.iv.2002, MA-02-13A-22"; 3[male] "3-11.iv.2002, MA-02-13A-23"; 2[male] "11-16.iv.2002, MA-02-13A24"; 2[male] "16-23.iv.2002, MA-02-13A-25"; 2[male] "23.iv-1.v.2002, MA-02-13A-26"; 3[male] "1-9.v.2002, MA-02-13A-27"; 4[male] "9-19.v.2002, MA-02-13A-28"; 3[male] "19-27.v.2002, MA-02-13A-29"; 2[male] "27.v-4. vi.2002, MA-02-13A-30"; 5[male] "4-12.vi.2002, MA-02-13A-31"; 1[male] "12-23.vi.2002, MA-02-13A-32"; 4[male] "3-13.vii.2002, MA-02-13A-34"; 1[male] "13-25.vii.2002, MA-02-13A-35"; 3[male] "8-19.viii.2002, MA-02-13A-37"; 4[male] "19.viii-3.ix.2002, MA-02-13A-38"; 1[male] "13-24.ix.2002, MA-02-13A-40"; 2[male] "24. ix-5.x.2002, MA-02-13A-41"; 1[male] "5-31.x.2002, MA-02-13A-42"; 1[male] "31.x-16.xi.2002, MA-02-13A43"; 2[male] "4-20.i.2003, MA-02-13A-47"; 4$ "MADAGASCAR: Tulear / Prov., Zombitse National / Park, near national road / 22[degrees]50.43'S, 44[degrees]43.87'E / 15.x-9.xi.2001, 825 m / R. Harin'Hala, Malaise / trap, deciduous spiny forest / MA-02-13B-01"; same except: 1[male] "9-19.xi.2001, MA-02-13B-02"; 1[male] "19-26. xi.2001, MA-02-13B-03"; 2[male] "26.xi-2.xii.2001, MA-02-13B-04"; 2[male] "7-14.xii.2001, MA-02-13B-06"; 7[male] "14-16.xii.2001, MA-02-13B-07"; 2[male] "22-29.xii.2001, MA-02-13B-09"; 7[male] "29.xii-5.i.2002, MA-02-13B- 10"; 1[male] "5-12.i.2002, MA-02-13B-11"; 3[male] "26.i-5.ii.2002, MA-02-13B-14"; 1[male] "5-14.ii.2002, MA-02-13B-15"; 4[male] "14-21.ii.2002, MA-02-13B-16"; 3[male] "28.ii-6.iii.2002, MA-02-13B-18"; 3[male] "6-13. iii.2002, MA-02-13B-19"; 3[male] "13-20.iii.2002, MA-02-13B-20"; 2[male] "20-27.iii.2002, MA-02-13B-21"; 1[male] "27.iii-3.iv.2002, MA-02-13B-22"; 1[male] "16-23.iv.2002, MA-02-13B-25"; 1[male] "19-27.v.2002, MA02-13B-29"; 2[male] "27-13.vii.2002, MA-02-13B-30"; 2[male] "13-25.vii.2002, MA-02-13B-31"; 1[male] "25. vii-1.viii.2002, MA-02-13B-32"; 1[male] "1-14.viii.2002, MA-02-13B-33"; 5$ "14-25.viii.2002, MA 02-13B-34 [1 with right wing detached and glued to card]"; 4[male] "25.viii-3.ix.2002, MA-02-13B-35"; 1[male] "MADAGASCAR: Tulear / Province, Beza Mahafaly / Reserve, Parcelle I near / research station / 23[degrees]41.19'S, 44[degrees]35.46'E / 15.x-10.xi.2001, 165 m / M.E. Irwin, F.D. Parker & / R. Harin'Hala, Malaise / trap in dry deciduous forest / MA-02-14A-01"; same except: 2[male] "28.xi-4.xii.2001, MA-02-14A-04"; 2$ "4-11.xii.2001, MA-02-14A-05"; 2[male] "11-18.xii.2001, MA-02-14A-06"; 2[male] "18-25.xii.2001, MA02-14A-07"; 1[male] "25.xii.2001-2.i.2002, MA-02-14A-08"; 3[male] "2-9.i.2002, MA-02-14A-09"; 1[male] "1618.i.2002, MA-02-14A-11"; 1[male] "25.i-1.ii.2002, MA-02-14A-13"; 2[male] "8-15.ii.2002, MA-02-14A-15"; 1[male] "22.ii-1.iii.2002, MA-02-14A-17"; 2$ "1-7.iii.2002, MA-02-14A-18"; 1[male] "14-22.iii.2002, MA-0214A-20"; 1[male] "28.vi-7.vii.2002, MA-02-14A-28"; 1[male] "18-28.vii.2002, MA-02-14A-30"; 1[male] "28.vii-9. viii.2002, MA-02-14A-31"; 1[male] "9-16.viii.2002, MA-02-14A-32"; 1[male] "16-28.viii.2002, MA-02-14A-33"; 2[male] "9-20.ix.2002, MA-02-14A-35"; 3[male] "20.ix-5.x.2002, MA-02-14A-36"; 1[male] "MADAGASCAR: Tulear / Province, Beza Mahafaly / Reserve, Parcelle II near / Bellevue, 23[degrees]41.39'S, 44[degrees]34.53'E / 15.x- 10. xi.2001, 180 m / M.E. Irwin, F.D. Parker & / R. Harin'Hala, Malaise trap / in spiny forest / MA-02-14B-01"; same except: 1[male] "4-11.xii.2001, MA-02-14B-05"; 1[male] "MADAGASCAR: Majunga / Ampijoroa National Park / 160 km N of Maevatanana / on RN 04 / 16[degrees]19.16'S, 46[degrees]48.80'E / 30.xi-8.xii.2003, 43 m / M.E. Irwin & R. Harin'Hala / Malaise trap in deciduous / Forest / MA-25-25"; 1[male] "MADAGASCAR: Tulear Province / Sous Prefecture Fort Dauphin / Andohaeala National Park / Ihazofotsy Parcelle III, 190 ft / 24[degrees]49.85'S, 46[degrees]32.17'E / 11-22.vi.2003 / Malaise trap, dry/spiny forest / M.E. Irwin, F. Parker & R. Harin'Hala / MG-21-22" (all CAS).

Other material examined (all labelled: "Curtonotum irwini sp. n. det. A.H. Kirk-Spriggs 2010"): MADAGASCAR: Tulear Prov.: 1[male] Zombitse National Park, near ANGAP office, 22[degrees]53.19'S:44[degrees]41.53'E, 21-28.ii.2002, 840 m, R. Harin'Hala, Malaise trap, deciduous spiny forest (MA-02-13A-17) [in spirit], BMSA(DNA)#0018; 1[male] same except: [in spirit], BMSA(DNA)#0019; 1[male] Beza Mahafaly Reserve, Parcelle I near research station, 23[degrees]41.19'S:44[degrees]35.46'E, 19.v-8.vi.2002, 165 m, R. Harin'Hala, Malaise trap in dry deciduous forest (MA-02-14A-25) [in spirit], BMSA(DNA)#0059. Majunga Prov.: 1[male] Ampijoroa National Park 160 km N of Maevatanana on RN 04, 16[degrees]19.16'S:46[degrees]48.80'E, 8-17.xii.2003, 43 m, M.E. Irwin & R. Harin'Hala, Malaise trap in deciduous forest (MA-25-26) [in spirit], BMSA(DNA)#0058 (all BMSA).

Distribution (Fig. 95): Occurring in the Western Dry Forest, South Western Dry Spiny Forest-Thicket, Wooded Grassland-Bushland and Western Sub-humid Forest vegetation types, in the Arid Spiny Bush and Dry Deciduous Forest biomes. In the Central and South biogeographical zones and Dry and Subarid bioclimatic zones (Figs 105-107; Tables 1-3; Appendix II).

Curtonotum keiseri Tsacas, 1974

Figs 1, 14, 27, 55, 58, 61, 79, 92

Curtonotum keiseri: Tsacas 1974: 706, 707 (figs 2e-f). Type locality: "Madagascar, Joffreville".

Differential diagnosis: This species is closely related to C. stuckenbergi Tsacas, differing in the colour of the frons (brown with distinct vittae in C. keiseri and yellow with indistinct vittae in C. stuckenbergi) the colour of flagellomere 1 and the shape of the male terminalia. Both share the deep brown, infuscate wing membrane, the dove-tailed sternite 6, and the straight, ventrally-directed, lateral spine and two smaller spines on the distiphallus. Curtonotum keiseri differ from C. stuckenbergi, however, in the angle and degree of curvature of the dm-cu crossvein of the wing, in the lateral margins of the phallus being only moderately sclerotised, and in the smaller spines of the distiphallus positioned in the basolateral region, rather than the left and right lateral regions. The ranges of the two species do not overlap, and they occur allopatrically. Redescription:

Male (primarily based on field-pinned HT and PT).

As redescribed for C. balachowskyi, differing in the following respects: Measurements: Total length 5 mm; length of head and thorax combined 3 mm; length of thorax and scutellum combined 3 mm (n = 1, PT); wing length 3.8 mm (n = 1, N-T).

Head (Figs 1, 14). As described for C. gladiiformis sp. n., except: eye height/length ratio: 13:8 (n = 1, PT); frons (Fig. 14), frons length/width ratio: 6:7 (n = 1, PT), orbital plates extending 0.9 length of frons; posterior orbital seta moderately strong, slightly shorter than outer vertical seta; flagellomere 1 yellow pruinose basally and along posterior margin, dark grey pruinose centrally and on anterior margin, arista with 10 or 11 long dorsal branches and 4 ventral branches in addition to terminal fork; gena narrow, eye height/genal height ratio: 12:1 (HT), silver pruinose, slightly darker beyond basal angle; vibrissae strongly developed; palpus pale brown.

Thorax (Fig. 1). Mesonotum with 2 median vittae wide, 2 lateral vittae shorter, clearly defined; supra-alar seta, ca twice length of posterior dorsocentral seta; postalar setae, moderately strong, slightly exceeding length of acrostichal setae; postpronotum with 13 finer, black-brown setulae; anepisternum surface with 33 fine setulae, some larger and arranged in 2 groups of 3 and 5; anepimeron, laterotergite and meron silver-grey to yellow-grey pruinose; katepisternum silver-grey to yellow-grey pruinose, dorsal katepisternal seta ca 0.3 length of ventral katepisternal seta, with 18 short, fine setulae at base and along posterior margin.

Scutellum. As described for C. gladiiformis sp. n.

Legs. Fore coxa with 13 brown setulae on anterior surface; mid coxa with 6 brown setulae; fore tibia with ctenidium of 12-14 short, sharp, black spinules.

Wing (as in Fig. 27). Long and broad, tip evenly-rounded, veins chestnut-brown, membrane deep-brown infuscate throughout, darker in r} and anterior half of r2+3 and in region of dm-cu crossvein; dm-cu crossvein with even arc dorsally; haltere dirty yellow.

Abdomen. Tergite 1 with oblique, small, subrectangular brown-black pruinose dorsolateral macula on either side and narrow medial facia; tergite 2 with larger subrectangular brown-black maculae and similar medial facia; tergites 3-5 with very wide, V-shaped concolourous median fascia and large concolourous T-shaped dorsolateral macula, these fully merging with medial fascia in anterior third, lateral margin of tergites 2-5 with subelliptical, large, concolourous macula in basal half; sternites 4-5 as described for C. coronaeformis sp. n.; sternite 6 (Fig. 79) dove-tailed (may appear narrower than Fig. 79 in undissected specimens), narrowed in basal third, with deep triangular excision apically, apical lobes evenly rounded, with dark brown maculae laterally, clothed in short, black, irregular, brown setulae in apical %, those at lateral and apical margins longer and more prominent.

Terminalia (Figs 55, 58, 61). Hypandrium (Fig. 55, hy) long, with broad-based rounded-truncate dorsobasal lobe, posterior bridge dorsally and ventrally produced (subtriangular to slightly angulate in profile); hypandrial arm constricted medially (viewed laterally), with 2 parallel setulae proximal to postgonite, of similar length (obscured by epandrium on Fig. 55), sclerotised area of medial lobes (viewed dorsally), with margins evenly rounded, convex medially, closely abutting, overlapping; postgonite (Fig. 55, pg) long, thin and straight, with slight undulating anterior margin; epandrium (Fig. 55, ep) slightly broader dorsally than ventrally (viewed laterally), evenly rounded on dorsal margin, posterior margin slightly angled, ventral margin with extensive row of long, regular to irregular, apically-directed setae; cercus (Fig. 55, ce) not prominent, longest setae longer than setae on dorsal margin of epandrium; surstylus (Fig. 55, ss) long, widest basally, slightly curved in apical %; phallus (as in Figs 58,ph, bp, dp, 61, bp, dp) C-shaped, moderately sclerotised, brown; phallapodeme (Fig. 58, ph) fused to basiphallus, subtriangular (viewed laterally), with basal margin developed into two flat, broad, subtriangular projections in basal fifth, bifurcated at point of connection with hypandrium; ejaculatory apodeme (Fig. 58, ea) free, duct inserted at junction of phallapodeme and basiphallus (missing from specimens illustrated in Fig. 58); basiphallus (Fig. 58, bp) broad basally and in region of first bend, then narrowed to apex, markedly narrowed in apical third (viewed dorsally); apical section (Figs 58, 61, bp) broad basally, sclerotised area extensive, abruptly narrowed towards apex, basal section with membranous window, with one narrow, but strong, straight, ventrally directed lateral spine (arrowed on Fig. 61) and two smaller spines, positioned in basolateral region, left margin of sclerotised area with irregular row of small tubules.

[FIGURES 55-63 OMITTED]

Variation: Insufficient material is available to assess variability.

Holotype (examined): [male] MADAGASCAR: "MADAGASCAR.D.-S. / Joffreville / 25.V.[19]58 F. KEISER [pink paper] // HOLOTYPE [red card] // CURTONOTUM / keiseri n.sp. / Holotype / L. TSACAS DET. 1973 [printed & handwritten] // BM [handwritten] // MUSEUM PARIS // Curtonotum / keiseri [male] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit 2006" [head missing] (MNHN). In poor condition, head and all legs except left fore femur, fore tibia, left hind femur and hind tibia missing; direct-pinned; dissected, abdomen and terminalia in micro-vial pinned beneath specimen.

Paratype: [male] "MADAGASCAR.D.-S. / Mtge. D'Ambre [= Montagne d'Ambre] / 24.V.[19]58 F.KEISER [pink paper] // PARATYPE [red card] // CURTONOTUM / keiseri n.sp. / paratype / L. TSACAS DET. 1973 [printed & handwritten] // Curtonotum / keiseri [male] / Tsacas, 1974 / A.H. Kirk-Spriggs vidit 2008" (NHMB).

Other material examined (labelled: "Curtonotum keiseri Tsacas, 1974 [male] det. A.H. Kirk-Spriggs 2011"): MADAGASCAR: 1[male] Madagascar N., Ambohitra, Joffreville, 800 m, 9-12.iv.1991, A. Freidberg & Fini Kaplan [left wing detached and glued to card] (TAU).

Distribution (Fig. 92): Apparently confined to the Humid Forest vegetation type in the Evergreen Rainforest biome. In the North biogeographical zone and Dry bioclimatic zone (Figs 105-107; Tables 1-3; Appendix II).
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Title Annotation:Part 1
Author:Kirk-Spriggs, Ashley H.
Publication:African Invertebrates
Article Type:Report
Geographic Code:6MADA
Date:Dec 1, 2011
Words:14079
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