Gilliesia ratchaburiensis , Boonsoong, Boonsatien & Sartori, Michel, 2015

Boonsoong, Boonsatien & Sartori, Michel, 2015, The nymph of Gilliesia Peters & Edmunds, 1970 (Ephemeroptera: Leptophlebiidae), with description of a new species from Thailand, Zootaxa 3981 (2), pp. 253-263: 254-263

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Gilliesia ratchaburiensis

sp. nov.

Gilliesia ratchaburiensis  sp. nov.

( Figs 1–40View FIGURE 1View FIGURES 2 – 5View FIGURES 6 – 9View FIGURES 10 – 13View FIGURES 14 – 22View FIGURES 23 – 29View FIGURES 30 – 34View FIGURES 35 – 38View FIGURES 39 – 40)

Material examined. HOLOTYPE: male imago (reared from nymph), THAILAND, Ratchaburi Province, tributary of Phachi River, 13 ° 28.6 ’ N, 99 ° 14.5 ’ E, 230 m, 30.iii. 2013, B. Boonsoong leg.. ALLOTYPE: female imago (reared from nymph), same data as holotype. PARATYPES: 4 mature nymphs, same data as holotype ( ZMKU); 4 nymphs mature nymphs, same data as holotype ( MZL); 9 nymph, 4.ii. 2013, same locale as holotype; 1 male imago (reared from nymph), 15.xi. 2014, same locale as holotype (all in ZMKU); 1 male imago (reared from nymph), 15.xi. 2014, same locale as holotype ( MZL).

Additional material. 7 nymphs, 2.ii. 2013, F. David leg.; 65 nymphs, 16.ii. 2013, F. David leg.; 6 nymphs, 30.ii. 2013, F. David leg.; same locale as holotype ( ZMKU).

Description. Mature nymph ( Fig. 1View FIGURE 1) (in alcohol). Body length between 3.0–4.0 mm for male (n= 10) and 3.5 – 5.0 mm for female (n= 10). Head prognathous, antennae more than 2 times width of head ( Fig. 1View FIGURE 1). Mouthparts ( Figs 2–16View FIGURES 2 – 5View FIGURES 6 – 9View FIGURES 10 – 13View FIGURES 14 – 22): Labrum with pronounced anteromedian emargination and row of setae on anterior margin directed ventrally ( Figs.2, 3View FIGURES 2 – 5, 14View FIGURES 14 – 22). Dorsal surface of labrum with scattered setae and no transverse row of setae. Ventral surface of labrum with distal row of bristles ( Figs. 4View FIGURES 2 – 5, 14View FIGURES 14 – 22). Left and right mandibles as in Figs 15–16View FIGURES 14 – 22, with well-developed prostheca, without seta on outer margin. Superlinguae of hypopharynx rounded, not extended laterally, with rows of setae along anterior margin; apex of lingua cleft ( Fig. 5View FIGURES 2 – 5); with two dense tufts of setae. Segment 1 of maxillary palpi subequal in length to segment 3. Length of segment 3 more than ca 1.6 times of length segment 2, apex of segment 3 with numerous long and thin setae, progressively longer from base to apex ( Figs 6–7View FIGURES 6 – 9). Maxillae with crown of setae covering inner 3 / 4 of apex of galea ( Figs 6–8View FIGURES 6 – 9); with 3 pectinate submarginal setae (ps, Figs 8–9View FIGURES 6 – 9), apex of maxilla with three reduced canines (c 1 –c 3, Fig. 9View FIGURES 6 – 9), proximal dentiseta (pd) and distal dentiseta (dd) entire ( Figs 8–9View FIGURES 6 – 9). Labium as in Figs 10–13View FIGURES 10 – 13; segment 2 of palpi slightly longer than 1 / 2 length of segment 1; segment 3 subequal in length to segment 2, triangular, with distinct stout bristles and thin setae ( Fig. 12View FIGURES 10 – 13); glossae distinctly ventral to paraglossae; narrow at top and wider at base, densely covered with long setae on ventral surface ( Fig. 13View FIGURES 10 – 13). Legs ( Figs 17–19View FIGURES 14 – 22): all legs with stout bristles on outer margin of femur and inner margin of tibia; tibio-patellar suture present on mid- and hindlegs; foreleg ( Fig. 17View FIGURES 14 – 22) with dark pigmentation on dorsal surface of femur, apex of tibia, and in middle of tarsi; midleg ( Fig. 18View FIGURES 14 – 22) with stout bristles on dorsal surface of femur; hindleg ( Fig. 19View FIGURES 14 – 22) with few short bristles on outer margin; tarsal claw ( Fig. 20View FIGURES 14 – 22) rather stout and not elongated, apex moderately hooked and narrow, with row of ca 10 teeth increasing in size towards apex. Abdominal terga with distinctive color pattern; terga 2–7 with the same pattern, with two median spots and two lateral spots; terga 2–9 with median longitudinal pale line; tergum 10 with two median spots ( Fig. 1View FIGURE 1). Posterolateral expansions present on abdominal segment 9 only ( Fig. 21View FIGURES 14 – 22). Gills ( Fig. 22View FIGURES 14 – 22): gills on segments 1–7 alike; long, slender and slightly forked at 2 / 5 distance from base; tracheae unbranched; no setae on gill margins. Caudal filaments with whorls of fine hair-like setae; terminal filament (3.4–3.8 mm) slightly longer than cerci (2.9–3.3 mm).

Male imago (holotype in alcohol). Body length 4.24 mm, fore wing 4.18 mm, hind wing 0.40 mm. Compound eyes black; antennae pale, pedicel brown, ocelli white with dark base ( Fig. 23View FIGURES 23 – 29). Vertex of head and notum brown. Forefemora ( Fig. 26View FIGURES 23 – 29) brown, with two white spots basally and medially; tibiae pale, brown at base and apex; tarsi pale, each segment distally brownish; ratios of segments in fore legs, 0.50: 1.00 (1.72 mm): 0.03: 0.37: 0.18: 0.15: 0.08. Mid- and hindlegs paler, with same markings as in foreleg. Midlegs ( Fig. 27View FIGURES 23 – 29): ratios of segments, 0.77: 1.00 (0.89 mm): 0.13: 0.05: 0.05: 0.10. Hind legs ( Fig. 28View FIGURES 23 – 29): ratios of segments, 0.78: 1.00 (0.93 mm): 0.11: 0.05: 0.05: 0.09. Forewing transparent, cross veins yellowish-brown; those in C-Sc, Sc-R 1 and R 1 - Rs surrounded by dark brownish clouds forming markings as shown in Fig. 24View FIGURES 23 – 29; MP 2 independent of vein MP 1; wing base brown; two long intercalaries between CuA and CuP ( Fig. 24View FIGURES 23 – 29). Hind wing very small with distinct acute costal projection near apex ( Fig. 25View FIGURES 23 – 29). Abdomen dark brown, terga 2–7 with one pair of median longitudinal white stripes arising from anterior margin; all terga with brown posterior margins; terga 8–9 entirely brown, tergum 10 with two pale spots ( Fig. 29View FIGURES 23 – 29). Caudal filaments missing. Genitalia ( Fig. 33View FIGURES 30 – 34): forceps length 0.4 mm (ratios of segments 1.00 (0.32 mm): 0.15: 0.09), slightly curved, first segment long and curved, second broader than third segment. Styliger plate with deep, V-shaped, median incision. Penis anchor-like, apical portion of each lobe bent laterally but not ventrally, sharply pointed, no spine arising from tip of penis lobe.

Female imago (allotype in alcohol). Body length 4.70 mm, fore wing 4.61 mm, hind wing 0.41 mm. Body color darker than in male. Vertex of head brown; compound eyes black; ocelli white with dark base ( Fig. 30View FIGURES 30 – 34). Pronotum pale, with dark brown pattern as in Fig. 30View FIGURES 30 – 34. Meso- and metanotum yellowish-brown, lateral and ventral parts of abdomen pale. All legs missing. Abdominal tergal patterns similar to that of male. Color pattern of wing similar to that of male but crossveins darker ( Fig. 31View FIGURES 30 – 34). Apex of sternum 9 with U-shaped, deep, median incision ( Fig. 34View FIGURES 30 – 34).

Egg. General shape ovoid; chorionic surface with small uniform tubercles subequal in size, each turbercle with indistinct ridges and shallow groove between turbercles ( Figs. 35–38View FIGURES 35 – 38).

Diagnosis and discussion. Male imagos of Gilliesia ratchaburiensis  n. sp. can be distinguished from those of G. hindustanica  and G. pulchra  by the following characters: 1) proportions of forelegs, 2) fore femur coloration, 3) abdominal terga patterns, 4) shape of apex of penis lobes, and 5) shape of hind wing ( Table 1).

abdominal terga dark brown with pitch brown translucent white on terga 1–8, dark brown on terga 1–10, with

on terga 1–8 with distinct markings distinct pale markings Nymphs of Gilliesia  can be distinguished from those of the two Leptophlebiinae  genera distributed in Southeast Asia ( Dipterophlebiodes  and Habrophlebiodes  ) by the posterolateral spines of the abdomen, gill shape, characteristics of the glossae, and maxillary palpi segment 3 ( Table 2). Gilliesia  seems to be more similar to Dipterophlebiodes  than to Habrophlebiodes  by the following characters: posterolateral spines well-developed on abdominal segment 9 only, and gills slender and slightly forked at> 1 / 3 distance from base ( Table 2).

Gilliesia  nymphs have all attributes of the subfamily Leptophlebiinae  as defined by Kluge (1994, 2009): 1) apex of maxilla with three reduced canines; proximal and distal dentisetae present and entire; 2) no transverse row of setae on the dorsal face of labrum; 3) tibio-patellar suture present on mid- and hindlegs; and 4) superlinguae of hypopharynx rounded, not extended laterally.

All these characters are considered by Kluge (1994) as plesiomorphic, hence the non-ranking plesiomorphon Leptophlebia  /fg 2 proposed by him ( Kluge 2009). Besides Gilliesia  , Leptophlebiinae  encompasses the following genera: Leptophlebia Westwood, 1940  (with three subgenera Leptophlebia  s.s., Paraleptophlebia Lestage, 1917  and Neoleptophlebia Kluge, 1997  ), Habrophlebiodes Ulmer, 1920  and Dipterophlebiodes Demoulin, 1954  . Many authors treat Paraleptophlebia  at the genus level, however.

Gilliesia  gills resemble somewhat those of Leptophlebia  s.s., being large in the proximal part before the fork, but contrary to Leptophlebia  s.s. the fork is not suddenly constricted to form a single filament but regularly decreasing, much more similar to what is found in Habroleptoides Schönemund, 1929 (Habrophlebiinae)  .

Gilliesia  tarsal claws are rather stout and not as elongated as in Leptophlebia  s.l. They are more similar to those of Dipterophlebiodes  or Habrophlebiodes  , the row of teeth extending over ¾ of the claw length, and teeth increasing slightly in size distally.

Habrophlebiodes  Dipterophlebiodes  Gilliesia 

posterolateral expansions of well developed on well developed on well developed on abdominal abdomen abdominal segments 8 and 9 abdominal segments 9 only segments 9 only

In Gilliesia  , the posterolateral spines are only developed on abdominal segment 9, as in Dipterophlebiodes  , whereas they are also present on segment 8 in Leptophlebia  s.l. (strongly developed in Leptophlebia  s.s. but more reduced in some Paraleptophlebia  ) and Habrophlebiodes  . In addition, maxillary canines of Gilliesia  are more reduced than in Habrophlebiodes  , ( Figs 39–40View FIGURES 39 – 40), whereas Peters (1972) did not mention the details of maxilla in Dipterophlebiodes  . The shape of Gilliesia  superlinguae resembles those of Dipterophlebiodes  , which are not extended laterally, whereas in Habrophlebiodes  they are slightly extended ( Peters & Edmunds 1970, Fig. 200). Gilliesia  possess two dense tufts of setae on ventral side of the lingua, as in Habrophlebiodes  ; this character is not mentioned for Dipterophlebiodes  ( Peters 1972, Fig. 5View FIGURES 2 – 5, p. 54).

At the imaginal stages, too, Gilliesia  is easily distinguished from Leptophlebia  s.l. by the presence of a hindwing costal process, and by the male penis lobes which do not possess “a ventral appendage arising from apex of each penis lobe” sensu Peters & Edmunds (1970, p. 177).

The eggs of Gilliesia  , here also described for the first time, resemble those of Habrophlebiodes  by the arrangement of chorionic tubercules (see Kang & Yang 1994, Fig. 17View FIGURES 14 – 22). Eggs of Leptophlebia  s.l. possess a very different chorionic structure (see for instance Koss 1968, Figs 30–36View FIGURES 30 – 34View FIGURES 35 – 38), without tubercles.

The genus Gilliesia  was previously known from northern India ( Gillies 1951, Peters & Edmunds 1970) and mainland China ( Zhou 2004). The discovery of a new species of this genus in Thailand extends its distribution to part of the Indo-Malayan mountain system in Southeast Asia, the locality of the new species being located in the Tenasserim Mountain Range. We may expect that some Oriental species described under the generic name Paraleptophlebia  at the nymphal stage only may actually be members of Gilliesia  . Among them, Paraleptophlebia spina Kang & Yang, 1994  from Taiwan could be a good candidate, because of the reduced maxillary canines ( Kang & Yang 1994, Fig. 8View FIGURES 6 – 9 D) and the shape of the eggs (ibid, Fig. 19View FIGURES 14 – 22).

Etymology. The specific epithet is named for the province, where the type material was collected: Ratchaburi province, Thailand.

Biological notes. The type-locality is located in a bamboo forest. The stream is characterized by a pronounced slope (25 % on the 200 m) and an alternation of riffles, pools and waterfalls up to 10 m high. The stream bed is mostly composed of boulders, cobbles, sand and leaf litter. The nymphs of Gilliesia ratchaburiensis  n. sp. were found in leaf bags of Bambusa bambos  and Lagerstroemia floribunda  in both pools and riffles habitats ( David & Boonsoong 2014). Nymphs were found mostly together with those of Habrophlebiodes prominens Ulmer, 1939  and Isca janiceae Peters & Edmunds, 1970  .

TABLE 1. Comparison of imaginal characteristics of three species of Gilliesia (Gillies 1951, Zhou 2004).

  tibiae equal in length to tarsi tibiae shorter than tarsi  
fore femur   dark brown with median pale

TABLE 2. Comparison of nymphal characteristics of three related genera (Habrophlebiodes, Dipterophlebiodes and Gilliesia).

  lingua rectangular or cordate with two dense tufts of setae, superlingua slightly extended laterally    
  length shorter than 1.6 times the length of segment 2; apical-pointed, with scattered setae length shorter than 1.6 times the length of segment 2; apical-blunted, with few setae  

Kiev Zoological Museum


Musee Zoologique














Gilliesia ratchaburiensis

Boonsoong, Boonsatien & Sartori, Michel 2015


Kluge 1997

Paraleptophlebia spina

Kang & Yang 1994


Demoulin 1954


Westwood 1940


Ulmer 1920


Lestage 1917