Corresponding author: Amir Yassin (
Academic editor: Rudolf Meier
A new classification of the subgenus
The drosophilid genus
In this paper, we propose a new classification based on recent phylogenetic findings, describe two new species, and provide a taxonomic key to all African
Frons and mesonotum of
Examined specimens were museum-preserved material or laboratory strains. Laboratory strains in the Laboratoire Evolution, Génomes et Spéciation (LEGS) belonged to 23 species (
List of laboratory strains used in studying internal reproductive structures and immature stages.
Species Founder | females collection data |
---|---|
Subgenus |
|
India: Bangalore; 2004, J. R. David | |
Subgenus |
|
Uganda: Kibale (1100 m); vii.2003, D. Lachaise | |
Tanzania: East Usambara Mt, Amani (900 m); 25-ix-2002, D. Lachaise | |
Tanzania: East Usambara Mt, Amani (900 m); 25-ix-2002, D. Lachaise | |
South Africa: Cape Town; ii.1984, J. R. David | |
Madagascar: Maroantsetra; 18-26.x.1987, S. F. McEvey, J. R. David & S. Aulard | |
Congo: Brazzaville; iii.2006, J. Vouidibio | |
Gabon: Ogoué-Ivindo, Makoukou (500 m); i.2004, F. Mavoungou | |
Congo: Brazzaville; iii.2006, J. Vouidibio | |
Brazil: Rio de Janeiro, Tijuca; 2001, J. R. David | |
Uganda: Kibale (1100 m); vii.2003, D. Lachaise | |
Madagascar: Antananarivo, Tsimbazaza (1200 m); ii.2008, A. Yassin & J. R. David | |
Tanzania: East Usambara Mt, Amani (900 m); 25-ix-2002, D. Lachaise | |
La Reunion (France): 2004, P. Capy | |
Madagascar: Andasibe; ii.2008, A. Yassin & J. R. David | |
Congo: Brazzaville; iii.2006, J. Vouidibiou | |
Kenya: S. Dupas | |
Sao Tomé & Príncipe: Pico de São Tomé Park (1500 m); iii.2001, D. Lachaise | |
Congo: Brazzaville; iii.2006, J. Vouidibiou | |
Kenya: S. Dupas | |
Sao Tomé & Príncipe: Pico de São Tomé Park (1500 m); iii.2001, D. Lachaise | |
Congo: Brazzaville; iii.2006, J. Vouidibiou | |
Madagascar: Antananarivo, Tsimbazaza (1200 m); ii.2008, A. Yassin & J. R. David | |
South Africa: Cape Province, Stellarbush; xii.2006, M. Debiais-Thibaud |
Formal morphological description of the new species followed standard
Morphological structures are abbreviated as: fw = front width; fl = front length; hw = head width; o = maximum diameter of the eye; j = width of gena in line with o; ch = maximum width of gena; or1 = proclinate orbital seta; or2 = anterior reclinate orbital seta; or3 = posterior reclinate orbital seta; oc = ocellar seta; poc = post-ocellar seta; iv = inner vertical seta; ov = outer vertical seta; acs = acrostichal setulae; adc = anterior dorsocentral; pdc = posterior dorsocentral; psc = prescutullar seta; bsc = basal scutellar seta; asc = apical scutellar seta; F1 = forefemur; WL = wing length; Wl = wing width; TL = thorax length; WV = width of white vittae at adc; BV = width of black vittae surrounding WV at adc; A = number of abdominal bristles summed over successive sternites. Measurements on immature stages were taken from uncrowded cultures grown under the same conditions (at 21°C). Measurements are abbreviated as: EL = egg length; El = egg width; PF = length of egg posterior filament; PL = puparium length; Pl = puparium width; H (horn index) = the ratio of the length of the anterior spiracles to the total length of the puparium × 100.
Mature, about 10 days old adults were dissected in a
1 | F1 without a row of spines ( |
2 |
– | F1 with a row of spines ( |
19 |
2(1) | F1 with a protruding tubercule bearing a bristle ( |
3 |
– | F1 without a protruding tubercule ( |
7 |
3(2) | Frons without a median white stripe; ♂A = 46–57; aedeagus subterminally concave ( |
|
– | Frons with a median white stripe; ♂A = 22–37; aedeagus subterminally convex ( |
4 |
4(3) | TST = 1–2 mm; spermatheca very papillate ( |
5 |
– | TST = 3–5 mm; spermatheca somewhat papillate ( |
6 |
5(4) | ♂WV = 1.5–1.8 μm; TST = 2.0 mm | |
– | ♂WV = 1.9–2.5 μm; TST = 1.2 mm | |
6(3) | TST = 3.2 mm | |
– | TST = 4.4 mm | |
7(2) | Frons without a median stripe | 8 |
– | Frons with a median stripe | 14 |
8(7) | Scutum velvety black, especially posteriorly; scutellum with a white spot at tip ( |
|
– | Scutum and scutellum not as above | 9 |
9(8) | Scutellum entirely and scutum posteromedially black ( |
|
– | Scutellum and scutum not as above | 10 |
10(9) | Wing darkened anteriorly | 11 |
– | Wing uniformally hyaline | 12 |
11(10) | Thorax and abdomen entirely dark brown ( |
|
– | Thorax and abdomen yellow ( |
|
12(10) | ♂ basitarsus without a hairy brush ( |
|
– | ♂ basitarsus with a hairy brush ( |
13 |
13(12) | Thorax yellow; the last 3 abdominal segments shining dark brown ( |
|
– | Thorax reddish yellow; abdomen shining yellow ( |
|
14(7) | Scutum with 6 longitudinal white stripes ( |
15 |
– | Scutum with 4 longitudinal white stripes ( |
16 |
15(14) | Aedeagal flap smooth and pointed basally ( |
|
– | Aedeagal flap finely serrated and truncated basally ( |
|
16(14) | Cercus with elongate, ventromedial expansion ( |
17 |
– | Cercus without ventromedial expansion ( |
18 |
17(16) | Thorax with a faint median white stripe ( |
|
– | Thorax without a faint median white stripe; ♂WL:TL = 2.25–2.35; abdomen without dark spots at the base of tergal bristles; cercal prominence short and almost entirely setulate along median edge ( |
|
18(16) | BV = 9–11 μm ( |
|
– | BV = 6–8 μm ( |
|
19(1) | F1 with spines not fused with long bristles at their bases ( |
20 |
– | F1 with spines fused with long bristles at their bases ( |
33 |
20(19) | F1 with 2 spines pointed in opposite orientation ( |
21 |
– | F1 with more than 2 spines usually pointed to the same direction ( |
22 |
21(20) | F1 small ( |
|
– | F1 large ( |
|
22(20) | ♂ basitarsus without a hairy brush | 23 |
– | ♂ basitarsus with a hairy brush | 24 |
23(22) | F1 with 3–4 spines; basalmost spine strong ( |
|
– | F1 with 5 spines internally and sometimes 2 spines externally ( |
|
24(22) | F1 spines differentiated; basalmost spine strong ( |
|
– | F1 spines undifferentiated ( |
25 |
25(24) | Wing anterior margin black or darkened ( |
26 |
– | Wing hyaline ( |
29 |
26(25) | Wing anterior margin black ( |
27 |
– | Wing anterior margin darkened ( |
28 |
27(26) | F1 with 2–3 spines ( |
|
– | F1 with 5–6 spines ( |
|
28(26) | F1 middle bristle borne on a tubercule ( |
|
– | F1 middle bristle not borne on a tubercule ( |
|
29(25) | Aedeagal flap absent ( |
30 |
– | Aedeagal flap present | 31 |
30(29) | F1 with a hairy tuft proximally ( |
|
– | F1 without a hairy tuft proximally ( |
|
31(29) | F1 middle bristle borne on a minute tubercule ( |
|
– | F1 middle bristle not borne on a tubercule ( |
32 |
32(31) | F1 not broadened, with a series of short bristles ( |
|
– | F1 broadened, with a few long bristles ( |
|
33(19) | WV < 15 μm; thorax and abdomen blackish brown | |
– | WV > 15 μm; thorax and abdomen not black | 34 |
34(33) | Abdominal tergal bristles with dark spots basally | 35 |
– | Abdominal tergal bristles without dark spots basally | 43 |
35(34) | Thorax with two incomplete submedian white stripes between two complete dorsocentral stripes ( |
|
– | Thorax without submedian stripes | 36 |
36(35) | F1 setiferous spines differentiated; basalmost borne on a protruding tubercule ( |
|
– | F1 setiferous spines undifferentiated | 37 |
37(36) | BV enlarged posteriorly; abdomen dark brown ( |
38 |
– | BV not enlarged posteriorly; abdomen light yellow | 40 |
38(37) | Abdomen darker than thorax ( |
|
– | Abdomen and thorax concolorous ( |
39 |
39(38) | First and second tarsomeres of the foreleg with strong black spines ( |
|
– | First and second tarsomeres of the foreleg without strong black spines; ♂TL = 1.44–1.56 mm ( |
|
40(37) | Head orange tan lighter than thorax ( |
|
– | Head and thorax concolorous reddish brown; hairy brush 2/3 ♂ basitarsus; spermatheca with an introvert ( |
41 |
41(40) | ♂ aedeagal flap highly serrated apically ( |
|
– | ♂ aedeagal flap highly smooth apically ( |
42 |
42(41) | ♂ aedeagal flap smooth basally ( |
|
– | ♂ aedeagal flap serrated basally ( |
|
43(34) | Abdomen yellow with brown posterior fine stripes on tergites II to IV; TST > 12.0 mm; spermatheca elongated ( |
|
– | Abdomen uniformally yellow; TST < 6.0 mm; spermatheca globulous (Figs 14b,c,e) | 44 |
44(43) | TST = 2.6 mm; spermatheca chitinized at base and apex ( |
|
– | TST = 4.0–5.2 mm; egg with 4 filaments | 45 |
45(44) | ♂TL = 1.56–1.70 mm; ♂ epandrium not expanded dorsally ( |
|
– | ♂TL = 1.44–1.50 mm; ♂ epandrium expanded dorsally ( |
Forefemur of
Classification and ecology of the subgenus Zaprionus. Breeding niches are abbreviated as: FL = flowers; FR = fruits; and TR = decaying tree trunk. Ability to be reared in the laboratory (L) is indicated as (+) for species that are reared and (-) for species that are not.
Group | Subgroup | Complex | Species | Authorship | L | Breeding niche | Reference |
---|---|---|---|---|---|---|---|
armatus | armatus | armatus | armatus | Collart, 1937a | - | TR: |
C37a |
FR: |
TC90 | ||||||
enoplomerus | Chassagnard, 1989 | + | FR: |
C89 | |||
seguyi | Tsacas & Chassagnard, 1990 | - | FR | TC90 | |||
spinipes | Tsacas & Chassagnard, 1990 | - | ? | TC90 | |||
spinoarmatus | Tsacas & Chassagnard, 1990 | - | FR: |
TC90 | |||
TR: |
B54 | ||||||
hoplophorus | hoplophorus | Tsacas & Chassagnard, 1990 | - | ? | |||
tuberarmatus | Tsacas & Chassagnard, 1990 | - | ? | ||||
vrydaghi | fumipennis | Séguy, 1938 | - | FL: |
TC90 | ||
vrydaghi | Collart, 1937a | - | FL: |
B76,C86 | |||
montanus | campestris | Chassagnard, 1989 | - | ? | C89 | ||
montanus | Collart, 1937b | - | TR: |
G57 | |||
TR: |
C89 | ||||||
spinosus | serratus | Chassagnard, 1989 | - | FL: |
C89 | ||
spineus | Tsacas & Chassagnard | - | ? | TC90 | |||
spinosus | Collart, 1937a | - | ? | ||||
inermis | arduus | Collart, 1937b | - | FR: |
C37b | ||
badyi | Burla, 1954 | - | FR | B54 | |||
ghesquierei | Collart, 1937a | - | FR: |
C37a | |||
FR: |
B54 | ||||||
FR: |
B76 | ||||||
FR: |
LT83 | ||||||
FR: |
L79 | ||||||
FR: |
L47 | ||||||
FR: |
L74 | ||||||
TR: |
L47 | ||||||
FR: |
C97 | ||||||
momorticus | Graber, 1957 | - | FL: |
G57 | |||
FL: |
L79 | ||||||
FL: |
L74 | ||||||
niabu | Burla, 1954 | - | FR: |
B54 | |||
inermis | cercus | Chassagnard & McEvey | + | FR: ex-banana trap | CM92 | ||
inermis | Collart, 1937a | + | FR: |
C37a | |||
FR: |
B54 | ||||||
TR: |
B54 | ||||||
FR: |
LT83 | ||||||
TR: |
LT83 | ||||||
tuberculatus | kolodkinae | Chassagnard & Tsacas, 1987 | + | FR: ex-banana trap | CT87 | ||
mascariensis | Tsacas & David, 1975 | + | FR: ex-banana trap | TD75 | |||
sepsoides | sepsoides | Duda, 1939 | + | FR: |
L79 | ||
FR: |
L82 | ||||||
FR: |
R83 | ||||||
FR: |
C86 | ||||||
tsacasi | Yassin, 2008 | + | FR: ex-banana trap | Y08 | |||
tuberculatus | burlai | Yassin, 2008 | + | FR: ex-banana trap | Y08 | ||
tuberculatus | Malloch, 1932 | + | FR: |
L79 | |||
FR: |
L82, LT83 | ||||||
FR: |
C86 | ||||||
verruca | Chassagnard & McEvey, 1992 | + | FR: ex-banana trap | CM92 | |||
neglectus | neglectus | Collart, 1937 | + | FR | B54 | ||
FL: |
B54 | ||||||
FL: |
C86 | ||||||
FR: |
L79 | ||||||
FR: |
C86 | ||||||
vittiger | davidi | davidi | Chassagnard & Tsacas, 1993 | + | FR: ex-banana trap | CT93 | |
taronus | Chassagnard & Tsacas, 1993 | + | FR: |
LT83 | |||
indianus | africanus | Yassin & David, 2008 | + | FR: ex-banana trap | Y08b | ||
gabonicus | Yassin & David, 2008 | + | FR: ex-banana trap | Y08b | |||
indianus | Gupta, 1970 | + | FR: ex-banana trap | Y08b | |||
FR: date palm, guava and citrus | Y09 | ||||||
ornatus | litos | Chassagnard & McEvey, 1992 | - | ? | CM92 | ||
ornatus | Séguy, 1933 | + | FR: |
B54 | |||
FR: |
C86 | ||||||
FR: |
L76 | ||||||
FR: |
L82, LT83 | ||||||
FL: |
L74 | ||||||
proximus | capensis | Chassagnard & Tsacas, 1993 | + | FR: ex-banana trap | CT93 | ||
proximus | Collart, 1937b | + | FR: |
C37b | |||
sexvittatus | multivittiger | Chassagnard, 1996 | + | FR: |
C96 | ||
sexstriatus | Chassagnard, 1996 | + | FR? | ||||
sexvittatus | Collart, 1937c | - | FR: |
C37c | |||
FR: |
C96 | ||||||
vittiger | camerounensis | Chassagnard & Tsacas, 1993 | + | FR: ex-banana trap | CT93 | ||
koroleu | Burla, 1954 | + | FR: ex-banana trap | ||||
TR: |
B54 | ||||||
lachaisei | sp. n. | + | FR: ex-banana trap | ||||
santomensis | sp. n. | + | FR: ex-banana trap | ||||
vittiger | Coquillett, 1902 | + | FR: ex-banana trap | YP |
References: B54:
It is still difficult to estimate with certainty the niches for some of the problematic species in
Distiphallus, testis and accessory gland, spermatheca and egg of
The armatus group was initially erected to include three subgroups: armatus, tuberculatus and vittiger (
Dorsal views of
The inermis group comprises species with spineless F1 (
Tarsomeres of male foreleg of
Ventral views of distiphallus of
We have recently collected a strain of
Lateral views of male epandrium and cercus and ventral views of aedeagus and hypandrium of
Ventral views of forefemur of
The vittiger group comprises 17 species with usually hairy epandrium carrying more than 2 posterior bristles (
Wing of
Lateral and dorsal views of
Puparium of
This species resembles
♂. TL = 1.38 mm.
Head. Arista with 3 dorsal and 2 ventral rays plus terminal fork; pedicel white, flagellomere I dark brown. Frons orange, without a median stripe but with orbital stripes inwardly bordered with black; ocellar triangle concolorous with frons; hw:fw = 2.42, fw:fl = 0.96. Orbital setae in straight line; or1:or2:or3 = 3:2:3, orbito-index = 1.1, oc:or1 = 1.45, poc:oc = 0.63, iv:ov = 0.88. Face whitish yellow; carina broad and bulbous. Gena broad, o:j = 9.3, o:ch = 6.2. Eye red.
Thorax. Scutum brown, darker than frons, with 2 silvery white stripes. acs in 6 rows in front of adc; adc:pdc = 0.8. Scutellum darker than scutum, with black borders of the stripes expanded posteriorly; bsc:asc = 0.7. Pleura yellow; sterno-index = 0.38. Forefemur with 4–5 spines borne on warts on the anteroventral margin. Male basitarsus with a hairy brush.
Wing. Yellowish. C-index = 2.5, 4v-index = 1.3, 4c-index = 0.9, 5x-index = 1.0, M-index = 0.4, ac-index = 2.5, b/c = 0.7, C3 fringe = 47%, and WL = 2.90 mm.
Abdomen. Entirely yellow with deep dark spots at the bases of tergal setae.
Terminalia. Epandrium densely pubescent throughout its entire length; posterior margin pubescent at dorsal portion with 4 long setae; epandrial ventral lobe with 3 long setae. Surstylus quadrate with two rows of prensisetae. Cercus triangular laterally. Hypandrium densely pubescent at the lateral portion of the paraphyses. Aedeagus expanded apically with a hook-like appendix; aedeagal flap expanded and deeply serrated. Apodeme subequal in length to aedeagus.
♀. TL = 1.39 mm, resembling male.
Terminalia. Oviscape constricted ventrally, with 8 peg-like and 6 short, marginal setae plus 4 supernumeraries. Spermatheca wide, campaniform and smooth.
Egg. Elliptical with 4, equally long and fine filaments.
Larva. Escaping the culture medium when crowded.
Puparium. Horn-index 9.8.
Distiphallus, oviscape and spermatheca of
This species resembles
♂. TL = 1.40 mm.
Head. Arista with 3 dorsal and 2 ventral rays plus terminal fork; pedicel white, flagellomere I dark brown. Frons orange, sometimes with highly vestigial median stripe plus orbital stripes inwardly bordered with black; ocellar triangle concolorous with frons; hw:fw = 2.45, fw:fl = 0.85. Orbital setae in straight line; or1:or2:or3 = 1.1:1.0:1.2, orbito-index = 1.1, oc:or1 = 1.4, poc:oc = 0.7, iv:ov = 0.7. Face whitish yellow; carina broad and bulbous. Gena narrow; o:j = 10, o:ch = 4.9. Eye red.
Thorax. Scutum brown, darker than frons, with 2 silvery white stripes. acs in 6 rows in front of adc; adc:pdc = 0.75. Scutellum darker than scutum, with black borders of the stripes expanded posteriorly; bsc:asc = 0.9. Pleura yellow; sterno-index = 0.44. Forefemur with 4–5 spines borne on warts on the anteroventral margin. Male basitarsus with a hairy brush.
Wing. Yellowish. C-index = 2.3, 4v-index = 1.4, 4c-index = 0.8, 5x-index = 1.0, M-index = 0.4, ac-index = 2.2, b/c = 0.6, C3 fringe = 52%, and WL = 2.7 mm.
Abdomen. Entirely yellow with deep dark spots at the bases of tergal setae.
Terminalia. Epandrium densely pubescent throughout its entire length; posterior margin pubescent at dorsal portion with 4 long setae; epandrial ventral lobe with 3 long setae. Surstylus quadrate with two rows of prensisetae. Cercus triangular laterally. Hypandrium densely pubescent at the lateral portion of the paraphyses. Aedeagus slender expanded apically without a hook-like appendix; aedeagal flap expanded and not serrated. Apodeme subequal in length to aedeagus.
♀. TL = 1.34 mm, resembling male.
Terminalia. Oviscape not constricted ventrally, with 6 (rarely 7) peg-like and 6 short, marginal setae plus 4 supernumeraries. Spermatheca globulous and smooth, not wider than longer.
Egg. Elliptical with 4 equally long and fine filaments.
Larva. Escaping the culture medium when crowded.
Puparium. Horn-index 10.4.
Male genitalia and spermatheca of
The identity of the dark species
Spermatheca and male epandrium of
Larval cephalopharyngeal skeleton of
This species resembles
♂. TL = 1.68 mm.
Head. Arista with 3 dorsal and 2 ventral rays plus a terminal fork, pedicel tan. Frons orange-tan with lateral white stripes; median white stripe absent; ocellar triangle raised and darker; hw:fw = 2.04, fw:fl = 1.05. Face pale; carina large; palpus yellow. Gena broad, o:j = 10.2, o:ch = 5.2. Orbital bristles in straight line; or2 very minute, or1:or2:or3 = 7:2:5, orbito-index = 1.4. Ocellar setae long, divergent; oc:or1 = 1.3, poc:oc = 0.5, iv:ov = 0.6. Eye red and densely pilose.
Thorax. Scutum tan, darker than frons, with four white longitudinal stripes continuing on scutellum; white stripes narrow, bordered with large black stripes, especially on the inner side; acs in 6 regular rows anterior to adc and 4 irregular rows between them; psc enlarged, adc:psc = 1.5; adc:pdc = 0.6. Scutellum slightly pointed at the apex, where white spot absent; bsc:asc = 1.3. Sterno-index = 0.6. F1 with 4 setiferous spines not borne on tubercules on the anteroventral margin. Basitarsus of the foreleg with a hairy brush on the ventral margin. Spiniform spines of the first and second tarsomeres of the foreleg enlaged and blackened.
Wing. Dusky; WL:WW = 2.3, C-index = 3.0, 4v-index = 1.5, 4c-index = 0.8, 5x-index = 0.7, M-index = 0.3, ac-index = 2.5, b/c = 0.6, C3 fringe 0.45, WL = 3.8 mm.
Abdomen. Uniformly tan, with dark spots at the bases of tergal bristles.
Terminalia (
♀. TL = 1.76 mm, resembling male.
Terminalia. Oviscape with 8 peg-like and 7 short, marginal setae plus 4 supernumary. Spermatheca large, globulous and smooth (
Egg. Elliptical with 4 equally long and fine filaments.
Larva. Not escaping the culture medium when disturbed or crowded.
Puparium. H = 5.0 (
. Tanzania.
. Holotype (male) and allotype (female), Tanzania: East-Usambara Mountains, Amani (870 m), ex type strain ZMI.12, 11-VIII-2008, founder female coll. 25-IX-2002, D. Lachaise. Paratypes: 10 males and 10 females with the same label. Types deposited in MNHN.
. Attempts to hybridize this strain with others belonging to the vittiger complex have all failed. The species is very prolific and easy to breed in the laboratory.
. Patronym, in honor of the French
This species resembles those of the indianus complex in having abdominal tergal spots and F1 spines not borne on protruding tubercule. It can be distinguished from them by the bigger body size, the darker body color mainly in contrast with the frons which is bright orange (
♂. TL = 1.40 mm.
Head. Arista with 2 dorsal and 3 ventral rays plus terminal fork; pedicel dark brown. Frons orange tan, with vestigial median stripe plus orbital stripes inwardly bordered with black; ocellar triangle blackened; hw:fw = 2.16, fw:fl = 0.8. Orbital setae in straight line; or1:or2:or3 = 3:2:3, orbito-index = 1.8, oc:or1 = 1.5, poc:oc=0.6, iv:ov = 0.4. Face tan. Gena narrow, o:j = 7.6, o:ch = 5.1. Eye red.
Thorax. Scutum brown, darker than frons, with 2 silvery white stripes. acs in 6 rows in front of adc; adc:pdc = 0.9. Scutellum darker than scutum, with black borders of the stripes expanded posteriorly; bsc:asc = 1.2. Pleura with white pilosity; sterno-index = 0.4. Forefemur with 4 spines not borne on warts on the anteroventral margin. Male basitarsus with a hairy brush.
Wing. Dusky; WL:WW = 2.3, C-index = 2.8, 4v-index = 1.4, 4c-index = 0.8, 5x-index = 0.9, M-index = 0.3, ac-index = 2.7, b/c = 0.6, C3 fringe 0.40, and WL = 3.2 mm.
Abdomen. Entirely yellowish, lighter than thorax, with faint dark spots at the bases of tergal setae.
Terminalia (
♀. TL = 1.50 mm, resembles male.
Terminalia. Oviscape with 8 peg-like and 6 short, marginal setae plus 4 supernumeraries. Spermatheca globulous and smooth (
Egg. Elliptical with 4 equally long and fine filaments.
Larva. Escaping the culture medium when crowded.
Puparium. Horn-index 10.6.
Sao Tomé and Príncipe.
Holotype (male) and allotype (female), Sao Tomé and Príncipe: Pico de São Tomé Park (1,500 m), ex type strain ZNG, 11-VIII-2008, founder female coll. III-2001, D. Lachaise. Paratypes: 10 males and 10 females with the same label. Types deposited in MNHN.
This species resembles
The species epithet is in reference to the type locality.
Many authors described the internal anatomy of some
Comparative morphometry of internal structures of male and female reproductive systems in
Male | Female | ||||||||
---|---|---|---|---|---|---|---|---|---|
TST | SV | VD | PAR | EC | EB | CAE | SR | SP | |
Subgenus |
|||||||||
|
4.4 | 2.0 | 0.80 | 2.6 | 2.2 | 0.30 | 0.6 | 3.8 | 0.07 |
Subgenus |
|||||||||
neglectus group | |||||||||
|
2.8 | 1.0 | 0.60 | 0.7 | 2.4 | 0.20 | 2.0 | 3.2 | 0.06 |
inermis group | |||||||||
|
1.2 | 0.6 | 0.04 | 2.0 | 1.1 | 0.22 | 1.0 | 1.5 | 0.04 |
|
1.5 | 1.1 | 0.20 | 2.6 | 2.1 | 0.32 | 0.4 | 1.0 | 0.09 |
|
1.4 | 0.9 | 0.16 | 2.2 | 2.0 | 0.22 | 1.6 | 0.9 | 0.08 |
|
4.4 | 0.9 | 0.40 | 3.2 | 1.1 | 0.22 | 0.5 | 7.2 | 0.12 |
|
1.0 | 0.7 | 0.20 | 1.6 | 2.1 | 0.20 | 0.8 | 0.8 | 0.06 |
|
2.0 | 0.6 | 0.20 | 3.2 | 1.6 | 0.20 | 0.1 | 1.0 | 0.04 |
|
1.3 | 0.8 | 0.40 | 3.6 | 1.2 | 0.20 | 0.4 | 1.2 | 0.06 |
|
3.2 | 1.2 | 0.70 | 2.2 | 0.9 | 0.20 | 0.3 | 3.6 | 0.06 |
|
4.4 | 1.0 | 1.10 | 2.0 | 1.3 | 0.12 | 0.3 | 6.3 | 0.06 |
|
3.8 | 1.6 | 0.80 | 2.0 | 2.0 | 0.20 | 1.2 | 4.0 | 0.06 |
vittiger group | |||||||||
|
12.4 | 7.2 | 2.20 | 3.6 | 0.9 | 0.30 | 0.7 | 12.0 | 0.18 |
|
5.3 | 2.2 | 1.30 | 2.2 | 1.5 | 0.30 | 0.7 | 4.8 | 0.16 |
|
5.4 | 1.0 | 0.70 | 1.6 | 1.3 | 0.30 | 0.8 | 3.8 | 0.07 |
|
2.5 | 0.7 | 0.40 | 0.7 | 0.7 | 0.16 | 0.4 | 3.5 | 0.06 |
|
2.6 | 1.4 | 0.80 | 2.0 | 1.6 | 0.30 | 0.6 | 3.0 | 0.06 |
|
5.2 | 1.4 | 1.40 | 3.2 | 2.2 | 0.30 | 0.8 | 4.6 | 0.06 |
|
4.0 | 2.0 | 0.80 | 2.6 | 1.2 | 0.30 | 0.6 | 4.6 | 0.07 |
|
3.6 | 2.4 | 2.00 | 1.4 | 2.0 | 0.28 | 0.3 | 4.2 | 0.06 |
3.6 | 1.6 | 1.20 | 2.0 | 1.6 | 0.34 | 0.7 | 3.2 | 0.10 | |
4.4 | 2.4 | 1.30 | 2.0 | 2.1 | 0.30 | 0.7 | 4.6 | 0.10 | |
|
4.4 | 2.4 | 1.30 | 2.0 | 2.4 | 0.30 | 0.8 | 4.2 | 0.12 |
|
4.2 | 2.0 | 0.70 | 3.2 | 1.2 | 0.20 | 0.6 | 4.5 | 0.09 |
TST = testis; SV = seminal vesicle; VD = vas deferens; PAR = paragonia (accessory gland); EC = ejaculatory bulb; CAE = caecum; SR = seminal receptacle; SP = spermatheca.
Measurements of immature stages in
Egg | Puparium | |||
---|---|---|---|---|
EL:El | PF:EL | PL:Pl | H | |
Subgenus |
||||
|
3.45 | 1.13 | 2.54 | 9.3 |
Subgenus |
||||
neglectus group | ||||
|
2.90 | 0.83 | 2.31 | 15.3 |
inermis group | ||||
|
3.00 | 0.54 | 2.54 | 9.4 |
|
3.26 | 1.13 | 2.62 | 13.1 |
|
2.90 | 0.97 | 2.40 | 10.3 |
|
2.91 | 0.73 | 2.47 | 6.8 |
|
2.75 | 0.97 | 2.43 | 9.0 |
|
3.10 | 0.90 | 2.57 | 8.6 |
|
2.73 | 0.90 | 2.53 | 8.4 |
|
2.86 | 0.90 | 2.59 | 7.0 |
|
3.00 | 0.91 | 2.29 | 7.2 |
|
3.40 | 0.88 | 2.31 | 10.6 |
vittiger group | ||||
|
3.18 | 1.14 | 2.52 | 10.0 |
|
3.44 | 0.81 | 2.49 | 8.3 |
|
3.26 | 0.90 | 2.46 | 9.8 |
|
3.33 | 0.83 | 2.43 | 10.4 |
|
3.05 | 1.16 | 2.54 | 10.5 |
|
2.87 | 0.91 | 2.29 | 12.0 |
|
2.43 | 1.00 | 2.45 | 9.8 |
|
3.67 | 1.06 | 2.44 | 10.6 |
2.86 | 0.60 | 2.24 | 10.6 | |
3.28 | 0.78 | 2.64 | 5.0 | |
|
3.20 | 1.06 | 2.65 | 9.3 |
|
3.00 | 0.93 | 2.56 | 11.0 |
EL = egg length; El = egg width; PL = puparium length; Pl = puparium width; H = horn-index.
Testis length (TST) ranges from 1.0 mm in
The seminal vesicle (SV) is the part of the vas deferens that has undergone a differentiation for sperm storage. It ranges from 0.6 mm in
The vas deferens (VD) ranges from 0.04 mm in
The ejaculatory bulb of
The seminal receptacle (SR) ranges from 0.8 mm in
The eggs of species of the Oriental subgenus
The length of the filaments varies between species (
Larvae of the genus
Puparia of the two subgenera are reddish brown in color (
The horn-index (H) is a classical taxonomic measurement in drosophilid systematics. H ranges from 5.0 in
Another important taxonomic character of the puparium is the branches of the anterior spiracle. In all
The two new species described here were collected by the late Dr. Daniel Lachaise (LEGS, CNRS) to which the authors would like to acknowledge their debt. The authors are also much grateful to two anonymous reviewers for constructive suggestions, Mrs. Marie-Thérèse Chassagnard and Dr. Léonidas Tsacas (MNHN) for allowing the reproduction of their drawings, Dr. Pierre Capy (LEGS) for providing the opportunity to travel to Madagascar from which several strains important to this study were collected, Dr. Shane F. McEvey (Australian Museum, Sydney) for his constructive critiques on a previous form of this manuscript, Dr. Brian Fischer (California Academy of Science), Dr. Lala Ravaomanarivo (University of Antananarivo) and workers in the Bibikely Biodiversity Institute for providing help during the collection of the flies, and to Drs. Vincent Debat (MNHN), Gerhard Baechli (Zoological Museum, Zurich), and Patrick Grootaert (Royal Belgian Institute of Natural Sciences, Brussels) for allowing the junior author to visit the collections and photograph the types. This work was funded by the Agence Universitaire de la Francophonie (AUF), Fondation des Treilles and Fondation Bettencourt-Schueller.