Caridina lanceolata Woltereck, 1937a

Rintelen, Kristina von & Cai, Yixiong, 2009, Radiation Of Endemic Species Flocks In Ancient Lakes: Systematic Revision Of The Freshwater Shrimp Caridina H. Milne Edwards, 1837 (Crustacea: Decapoda: Atyidae) From The Ancient Lakes Of Sulawesi, Indonesia, With The Description Of Eight New Species, Raffles Bulletin of Zoology 57 (2), pp. 343-452 : 369-373

Caridina lanceolata Woltereck, 1937a

( Figs. 13-14 , Table 6)

Caridina lanceolata Woltereck, 1937a: 224 , Figs. I.7 a-c, pls. 3,6 (type locality: Lake Towuti at Lingkona, Lake Mahalone [Mahalona], Lake Matanno [Matano] at Bonti, Lake Matanno at Soroako).

Caridina lanceolata – Woltereck, 1937b: 307, fig. 11; Chace, 1997: 12; Fernandez-Leborans et al., 2006a: 1, fig. 24; 2006b: 1985, Table I; Roy et al., 2006: 1090; von Rintelen et al., 2007b: 262, fig. 2b; 2007b: 4, fig. 2, 2008: 2244, Figs. 1 , 4 , Tables 1, 2; Cai et al., 2009: 15, Figs. 1-2 (type locality of neotype: Lake Matano, Mengonuwai).

Cardina lanceolata – Brooks, 1950: 168 (erroneous spelling).

Material examined. – Lake Matano: 10 ex. ( ZMB 29021, some SEM material), north shore, 02°29.75'S, 121°25.81'E, loc. 42- 03, coll. K . & T. von Rintelen , 17 Sep.2003 ; 112 ex. ( MZB Cru 1738, n=55; ZMB 29070, n=57), south shore, at small islands, 02°28.458'S, 121°15.57'E, loc. F4-02, mixed substrate, coll. F. Herder, 2 Nov.2002 ; 5 ex. ( ZMB 29082), northwest corner, 02°27.71'S, 121°13.03'E, loc. 102-03, on rocks in deeper water, coll. K . & T. von Rintelen , 7 Oct.2003 ; north shore, 02°26.36'S, 121°19.03'E, loc. 84-03, on wood ( ZMB 29090, n=3), coll. K . & T. von Rintelen , 1 Oct.2003 ; 21 ex. ( MZB Cru 1739, n=10, ZMB 29110; n=11), south shore, near cave entrance, 02°29.85'S, 121°18.66'E, loc. 60-03, on rocks in deeper water, coll. K . & T. von Rintelen , 24 Sep.2003 ; 31 ex. ( MZB Cru 1740, n=14, ZMB 29111, n=17), south shore, west of small islands, 02°28.5'S, 121°15.55'E, loc. 82-03, on macrophytes, coll. K . & T. von Rintelen , 1 Oct.2003 ; 129 ex. ( MZB Cru 1741a, n=35; ZMB 29120a, n=10 and MZB Cru 1742b, n=50; ZMB 29120b, n=34, some SEM material), north cl (mm) 2.2-4.0 3.0 ± 0.4 3.1 129 rl / cl 1.5-2.3 1.9 ± 0.2 1.9 120 n dorsal rostral teeth 8-19 14 ± 2 13 124 n ventral rostral teeth 4-13 7 ± 2 7 124 abds6 / cl 0.8-1.1 0.9 ± 0.0 1.0 129 abds6 / abds5 1.4-2.3 2.0 ± 0.1 2.0 33 abds6 / h tel 1.0-1.9 1.3 ± 0.1 1.3 101 h tel / w tel 3.3-3.7 3.5 ± 0.1 3.5 6 n spines uropodal diaeresis 7-9 8 ± 1 8 5 h ch1 / w ch1 1.9-3.5 2.6 ± 0.4 2.6 35 h ch1 / h ca1 0.8-1.4 1.1 ± 0.1 1.1 127 h ca1 / w ca1 2.4-5.3 3.6 ± 0.7 3.7 35 h ch2 / w ch2 2.1-6.7 3.7 ± 0.8 3.7 35 h ch2 / h ca2 0.5-1.6 0.7 ± 0.1 0.6 127 h ca2 / w ca2 5.4-9.9 8.3 ± 1.2 8.6 35 n spines p3 4-6 5 ± 1 6 5 n spines p5 28-35 31 ± 3 31 5

shore, 02°28.42'S, 121°24.21'E, loc. 41-03, (a) on wood, (b) on macrophytes, coll. K. & T. von Rintelen, 17 Sep.2003; 2 ex. ( ZMB 29160) , west shore, 02°26.828'S, 121°12.988'E., loc. 135-04, on wood, coll. K. & T. von Rintelen & A. Wessel, 23 Jul.2004; 20 ex. ( MZB Cru 1743, n=11; ZMB 29162, n=9) , south shore, 02°27.84'S, 121°13.88'E, loc. 63-03, on mixed substrate, coll. K. & T. von Rintelen, 24 Sep.2003; 26 ex. ( MZB Cru 1744, n=13; ZMB 29164, n=13, some SEM material) , east shore, outlet bay, at Petea outlet, 02°32.11'S, 121°28.69'E, loc. 44-03, on macrophytes, coll. K. & T. von Rintelen, 18 Sep.2003; 40 ex. ( MZB Cru 1745, n=20; ZMB 29190, n=20) , north shore, 02°27.28'S, 121°21.21'E, loc. 98-03, on mixed substrate, coll. K. & T. von Rintelen, 5 Oct.2003; 2 ex. ( ZMB 29193) , east shore, just south of entrance to outlet bay, 02°31.54'S, 121°27.0'E, loc. 43-03, on rocks in shallow water, coll. K. & T. von Rintelen, 18 Sep.2003; 2 ex. ( ZMB 29194) , southwest shore, at Cape Nikomene , 02°32.24'S, 121°24.76'E, loc. 46-03, on rocks, coll. K. & T. von Rintelen, 18 Sep.2003; 7 ex. ( ZMB 29237) , north shore, 02°25.67'S, 121°16.54'E, loc. 65-03, on leaf litter, coll. K. & T. von Rintelen, 25 Sep.2003; 18 ex. ( ZMB 29249) , south shore, canal between island and mainland, 02°28.46'S, 121°15.83'E, loc. 62-03, on mixed substrate, coll. K. & T. von Rintelen, 1 Oct.2003; 14 ex. ( ZMB 29321) , south shore, 02°28.461'S, 121°15.591'E, loc. 137-04, on macrophytes, coll. K. & T. von Rintelen, 24 Sep.2004; 2 ex. ( ZMB 29322) , south shore, at small islands, 02°28.476'S, 121°15.64'E, loc. 138-04, on wood, coll. K. & T. von Rintelen & A. Wessel, 24 Jul.2004; 9 ex. ( ZMB 29323) , south shore, 02°27.85'S, 121°13.87'E., loc. 125-04, on rocks in deeper water, coll. P. Koller & K. von Rintelen, 1 Aug.2004; 1 ex. ( MZB Cru 1746) , east shore, south bay, 02°32.77'S, 121°26.71'E, loc. 45-03, on rocks, coll. K. & T. von Rintelen, 18 Sep.2003; 1 ex. ( ZMB 29188) , Petea River , 02°32.64'S, 121°29.51'E, loc. 101-03, on leaf litter, coll. K. & T. von Rintelen, 6 Oct.2003; 15 ex. ( MZB Cru 1762, n=8; ZMB 29279, n=7) , Petea River , 02°32.672'S, 121°30.137'E, loc. F4-04, substrate unknown, coll. F. Herder, 25 Mar.2004 .

Lake Mahalona – 41 ex. ( MZB Cru 1747a, n=2; ZMB 29022a, n=21 and MZB Cru 1748b, n=9; ZMB 29022b, n=9, some SEM material), northwest shore, 02°34.72'S, 121°29.12'E, loc. 56-03, on (a) on wood, (b) on macrophytes, coll. K . & T. von Rintelen , 23 Sep.2003 ; 22 ex. ( MZB Cru 1749, n= 11; ZMB 29041, n=11), east shore, 02°34.54'S, 121°30.48'E, loc. 55-03, on Ottelia , coll. K . & T. von Rintelen , 23 Sep.2003 ; 12 ex. ( MZB Cru 1750, n=6; ZMB 29042, n=6, some SEM material), south shore, 02°36.93'S, 121°30.02'E, loc. 57-03, on Ottelia , coll. K . & T. von Rintelen , 23 Sep.2003 ; 18 ex. ( MZB Cru 1751, n=9; ZMB 29236; n=9, some SEM material), east shore, 02°34.217'S, 121°30.681'E, loc. 147-04, on macrophytes, coll. K . & T. von Rintelen , 3 Aug.2004 ; 31 ex. ( MZB Cru 1752, n=15; ZMB 29250, n=16), south shore, 02°36.982'S, 121°30.078'E, loc. 45-04, on wood, coll. M. Glaubrecht & T. von Rintelen , 26 Mar.2004 ; 1 ex. ( ZMB 29336), north shore, at cape, 02°34.71'S, 121°29.144'E, loc. 148-04, on mixed substrate, coll. K . & T. von Rintelen , 3 Aug.2004 ; 15 ex. ( MZB Cru 1753), east shore, estuary of Ponsoa River, 02°35.101'S, 121°30.681'E, loc. 146-04, on Ottelia , coll. P. Koller, K . & T. von Rintelen , 3 Aug.2004 ; 8 ex. ( ZMB 29115), Tominanga River , approx. 2.2 km north of Lake Towuti, 02°36.5'S, 121°31.78'E, loc. 58-03, on macrophytes, coll. K . & T. von Rintelen , 23 Sep.2003 .

Lake Towuti – 69 ex. ( MZB Cru 1754, n=59; ZMB 29029, n=10, some SEM material), south shore, approx. 100 m offshore, 02°56.27'S, 121°23.67'E, loc. 75-03, on Ottelia , coll. K . & T. von Rintelen, 28 Sep.2003 ; 2 ex. ( ZMB 29030), west shore, 02°48.526'S, 121°25.044'E, loc. 120-04, on wood, coll. K . & T. von Rintelen & A. Wessel, 29 Jul.2004 ; 141 ex. ( MZB Cru 1755, n=123; ZMB 29032, n=18, some SEM material), west shore, outlet bay, east of Cape Kombe , 02°48.08'S, 121°23.05'E, loc. 53-03, on mixed substrate, coll. K . & T. von Rintelen, 21 Sep.2003 ; 34 ex. ( MZB Cru 1756, n=24; ZMB 29033a, n=8 and ZMB 29033b, n=2, some SEM material), southwest shore, Cape Sioloya , 02°50.7'S, 121°26.32'E, loc. 77-03, ( MZB and ZMB a) pelagic, (b) on wood, coll. K . & T. von Rintelen, 28 Sep.2003 ; 3 ex. ( ZMB 29081), east shore, south of Cape Tomeraka , 02°44.47'S, 121°37.53'E, loc. 70-03, on rocks, coll. K . & T. von Rintelen, 9 Oct.2003 ; 206 ex. ( MZB Cru 1757a, n=22; ZMB 29093a, n=11 and MZB Cru 1758b, n=92; ZMB 29093b, n=81, some SEM material), southwest shore, west of Cape Tetetu , 02°454.13'S, 121°23.78'E, loc. 76-03, (a) on wood, (b) pelagic, coll. K . & T. von Rintelen, 28 Sep.2003 ; 10 ex. ( ZMB 29094, some SEM material), northeast shore, at Cape Noote , 02°39.751'S, 121°39.195'E, loc. 117-04, on wood, coll. K . & T. von Rintelen, 28 Jul.2004 ; 7 ex. ( ZMB 29112), Loeha Island , south shore, 02°46.85'S, 121°32.86'E, loc. 96-03, on macrophytes, coll. K . & T. von Rintelen, 4 Oct.2003 ; 9 ex. ( ZMB 29113), east shore, off Beau village , 02°48.99'S, 121°33.64'E, loc. 71-03, on Ottelia , coll. K . & T. von Rintelen, 27 Sep.2003 ; 55 ex. ( MZB Cru 1759, n=27; ZMB 29114, n=28), west shore, outlet bay, west of Cape Tokaluku , 02°47.32'S, 121°23.38'E, loc. 52-03, on macrophytes, coll. K . & T. von Rintelen, 21 Sep.2003 ; 1 ex. ( ZMB 29128), west shore, at entrance to outlet bay, Cape Larona , 02°48.43'S, 121°24.75'E, loc. 73-03, on wood, coll. K . & T. von Rintelen, 27 Sep.2003; ( ZMB 29158, few juveniles) , north shore, west of Cape Manu , 02°41.67'S, 121°36.85'E, loc. 69-03, on macrophytes, coll. K . & T. von Rintelen, 27 Sep.2003 ; 4 ex. ( ZMB 29161), Larona River , close to outlet bay, 02°45.8'S, 121°20.8'E, loc. 51-03, on macrophytes, coll. K . & T. von Rintelen, 21 Sep.2003 ; 13 ex. ( MZB Cru 1761, n=6; ZMB 29163, n=7), Larona River , close to outlet bay, 02°45.6'S, 121°20.12'E, loc. 50-03, on macrophytes, coll. K . & T. von Rintelen, 20 Sep.2003; ( ZMB 29166, few juveniles) , east shore, 02°52.79'S, 121°31.18'E, loc. 72-03, on Ottelia , coll. K. & T. von Rintelen, 27 Sep.2003 ; 7 ex. ( ZMB 29167), Loeha Island , south shore, 02°47.22'S, 121°33.68'E, loc. 91-03, on Ottelia , coll. K . & T. von Rintelen, 2 Oct.2003 ; 12 ex. ( ZMB 29181), Loeha Island , west shore, 02°45.5'S, 121°31.06'E, loc. 95-03, on rocks, coll. K . & T. von Rintelen, 4 Oct.2003 ; 23 ex. ( ZMB 29182, some SEM material), north shore, bay east of Cape Bintu , 02°39.48'S, 121°33.25'E, loc. 68-03, on rocks, coll. K . & T. von Rintelen, 26 Sep.2003; ( ZMB 29183, few juveniles) west shore, north of Cape Wasupute , 02°46.9'S, 121°27.94'E, loc. 78-03, on rocks, coll. K . & T. von Rintelen, 28 Sep.2003 ; 4 ex. ( ZMB 29314), west shore, outlet bay, 02°47.142'S, 121°23.63'E, loc. 11-05, on mixed substrate, coll. K . & T. von Rintelen, 9 Jan.2005 ; 4 ex. ( ZMB 29315), west shore, Cape Bakara , 02°40.771'S, 121°26.11'E, loc. 144-04, on mixed substrate, coll. K . & T. von Rintelen, 26 Jul.2004 ; 12 ex. ( ZMB 29317), west shore, west of Cape Timbalo , 02°42.631'S, 121°26.389'E, loc. 145-04, on wood, coll. K . & T. von Rintelen, 26 Jul.2004 ; 4 ex. ( ZMB 29318), northwest shore, south of Timampu , 02°39.466'S, 121°25.859'E, loc. 140-04, on macrophytes, coll. K . & T. von Rintelen, 25 Jul.2004 ; 43 ex. ( MZB Cru 1760, n= 28; ZMB 29319, n=15), outlet bay, at Cape Kombe , 02°48.083'S, 121°23.049'E, loc. 118-04, ( MZB) on macrophytes, ( ZMB) on wood, coll. K . & T. von Rintelen, 29 Jul.2004 ; 8 ex. ( ZMB 29320), northwest shore, 02°40.897'S, 121°25.015'E, loc. 143-04, on wood ( Pandanus ), coll. K . & T. von Rintelen, 25 Jul.2004 .

Description. – Carapace length 2.2-4.0 mm (n=129). Rostrum ( Fig. 14 A-C; Table 6) very long, slender and always strongly upturned, reaching far beyond end of scaphocerite, 1.5-2.3 times as long as carapace (n=120), armed dorsally with 8- 19 teeth (including 1-3 teeth posterior to orbital margin), approx. anterior third to half unarmed, usually with 1-4 subapical teeth, sometimes subapical teeth absent, armed ventrally with 4-13 teeth (n=124). Antennal spine situated below inferior orbital angle. Pterygostomial angle broadly rounded. Eyes well developed, anterior end 0.6-0.7 times length of basal segment of antennular peduncle (n=5). Antennular peduncle 0.9-1.0 times as long as carapace (n=5), second segment 2.0-2.4 times length of third segment, third segment 0.2-0.3 times length of basal segment. Stylocerite reaching 0.87-0.9 times length of basal segment of antennular peduncle (n=5). Scaphocerite ( Fig. 14F ) 4.8-5.4 times as long as wide (n=5).

Sixth abdominal somite 0.8-1.1 times length of carapace (n=129), 1.4-2.3 times as long as fifth somite (n=33), 1.0- 1.9 times length of telson (n=101). Telson ( Fig. 14G,J ) 3.3-3.7 times as long as wide (n=6), distal margin rounded, without projection, with 3-5 pairs of spinules and 1 pair of dorsolateral spinules; distal end with 4 pairs of spines, lateral pair distinctly longer than intermediate pairs, median pair shortest. Preanal carina ( Fig. 14E ) with a spine. Uropodal diaeresis ( Fig. 14D ) with 7-9 movable spinules (n=5).

5 pairs of pleurobranchs well developed; 3 pairs of arthrobranchs, 2 on third maxillipeds, with second pair strongly reduced in size, 1 pair on first pereiopod; 1 pair of podobranchs on second arthrobrach maxillipeds reduced strongly to a laminate form. Epipods only present on first two pereiopods. Mouthparts as described by Cai et al. (2009).

Chela and carpus of first pereiopod distinctly stouter and broader than chela and carpus of second pereiopod ( Fig. 14 M-O); chela of first pereiopod 1.9-3.5 times as long as wide (n=35), 0.8-1.4 times length of carpus (n=127); tips of fingers rounded, without hooks; dactylus 1.2-1.5 times as long as palm (n=7); carpus 2.4-5.3 times as long as wide (n=35), 1.2-1.4 times length of merus (n=5). Chela of second pereiopod 2.1-6.7 times as long as wide (n=35), 0.5-1.6 times length of carpus (n=127); tips of fingers rounded, without hooks, dactylus 1.2-1.7 times as long as palm (n=7); carpus 5.4-9.9 times as long as wide (n=35), 1.5-1.6 times as long as merus (n=5).

Third pereiopod ( Fig. 14H,K ) slender, dactylus 4.5-5.4 times as long as wide (terminal spine included, without spines of flexor margin; n=5), terminating in one large claw with 4-6 accessory spines on flexor margin; propodus 12.3-18.8 times as long as wide, 3.4-4.5 times as long as dactylus; carpus 5.8-6.8 times as long as wide, 0.5 times as long as propodus, 0.5 times as long as merus; merus 10.5-11.4 times as long as wide, bearing 3-4 strong, movable spines on posterior margin of outer surface.

Fifth pereiopod slender ( Fig. 14I,L ), dactylus 4.0-6.2 times as long as wide (terminal spine included, without spines of flexor margin; n=5), terminating in one large claw with 28-35 accessory spines on flexor margin; propodus 11.3-21.3 times as long as wide (n=5), 3.3-5.3 times as long as dactylus; carpus 5.0-6.3 times as long as wide, 0.5 times as long as propodus, 0.5-0.6 times as long as merus; merus 8.3-11.5 times as long as wide, bearing 2-3 strong, movable spines on posterior margin of outer surface.

Endopod of male first pleopod ( Fig. 14P ) elongated triangular, 1.4-2.5 times as long as proximally wide (n=5), without appendix interna. Appendix interna of male second pleopod ( Fig. 14Q ) 0.9-1.0 times length of appendix masculina (n=5).

Ovigerous females with 15- 69 eggs (n= 27 females); egg size 0.6-0.9 x 0.4-0.6 mm (n=701, eggs with and without eyes).

Distribution. – Endemic to the Malili lake system, widely distributed and abundant in Lake Matano, Lake Mahalona, Lake Towuti, and connecting rivers, i.e. Larona (close to the outlet of Lake Towuti), Petea, and Tominanga ( Fig. 13A ). As these rivers are rather an extension of the lakes themselves (albeit with currents), C. lanceolata is regarded here, in contrast to C. masapi , as a typical lacustrine species.

Biology and ecology. – C. lanceolata occurs on various kinds of substrates, except sponges, that are available in the Malili lakes (rocks from gravel to boulders, wood, leaf litter, different kinds of macrophytes, and in pelagic swarms), and down to a depth of approx. 10 m. When disturbed, C. lanceolata makes characteristic abrupt movements in all directions or simply keeps still in midwater. It often occurs syntopically, i.e. on the same substrate, with other species, for example with C. tenuirostris on wood in Lake Towuti and Lake Mahalona. It is the only wood dweller in Lake Matano and often the only shrimp on macrophytes surrounded by sand or mud, e.g. on the basal parts of the water plant Ottelia .

Colour pattern. – Body and appendages are transparent throughout, with a reddish pigmentation caused by several red, sometimes yellowish-greenish, dots or spots ( Fig. 13B ). Ovigerous females were observed to bear green eggs.

Taxonomic remarks. – Cai et al. (2009: 19) designated a neotpye for C. lanceolata : “Specimens of Woltereck (1937a, b) are no longer extant […]. As all new species described by Woltereck are morphologically close to each other and to stabilize the taxonomic status of those species, neotypes are designated if specimens are available from the recent collections. A neotype from Lake Towuti is designated here (1 male, cl 2.9 mm, ZRC, Lake Matano, south coast)” .

C. lanceolata can easily be distinguished from all other ancient lake species by the characteristic shape and denticulation of the rostrum, which is already visible in juveniles: strongly upturned with approximately anterior third to half unarmed and usually with 1-4 subapical teeth. The arrangement of rostral teeth slightly resembles C. tenuirostris , but this species has a distinctly more slender rostrum, a higher number of ventral teeth (10-24, median 15 vs. 4-13, median 7 in C. lanceolata ), and subapical teeth are always absent. Also, C. lanceolata differs from all other species by a distinctly longer sixth abdominal somite compared to the carapace length (0.8-1.1, median 1.0 vs. shorter in all other species).

Roy et al. (2006) hint at the possibility that C. lanceolata might comprise cryptic species, but a recent study by von Rintelen et al. (in review) does not support this hypothesis. Although there are genetic differences between allopatric populations from the three major lakes of the Malili system, C. lanceolata is here regarded as a valid taxon without the need of further taxonomic subdivision. C. lanceolata is genetically distinct from all other ancient lakes species ( Figs. 63-64 ), and the clade only consisting of C. lanceolata specimens even represents a single colonization event of the Malili lakes ( Fig. 63 ).

Figures

Fig. 13. Caridina lanceolata from the Malili lake system. A. Distribution. B. Colour pattern of living animal (not to scale). Picture courtesy of Chris Lukhaup.

Fig. 14. Caridina lanceolata from the Malili lake system. A. Cephalothorax and cephalic appendages, female (ZMB 29032); B. Rostrum variability (ZMB 29033a); C. Woltereck’s drawings of the rostrum, from several lakes (modified from 1937a); D. Uropodal diaeresis, female (ZMB 29032); E. Preanal carina, male (ZMB 29093a), F. Scaphocerite; G. Telson, female (ZMB 29032); H. Third pereiopod, female (ZMB 29093b); I. Fifth pereiopod; J. Distal end of telson, female (ZMB 29032); K. Dactylus of third pereiopod, female (ZMB 29093b); L. Dactylus of fifth; M. first pereiopod, female (ZMB 29032); N. Second pereiopod; O. SEM image of chela and carpus of first and second pereiopods; P. Endopod of male first pleopod (ZMB 29093a); Q. Appendix masculina of male second pleopod. Scale bars: A-B, F = 1.0 mm; E, G-I, M-Q = 0.5 mm; D, J-L = 0.1 mm; C = no scale available.

Fig. 7. Caridina glaubrechti from the Malili lake system. A. Distribution. B. Colour pattern of living animal (not to scale). Picture courtesy of Chris Lukhaup.

Fig. 1. The ancient lakes of Sulawesi. A. The two ancient lake systems on the Indonesian island Sulawesi. B. Overview of Caridina collecting sites from Lake Poso. C. Overview of Caridina collecting sites from the Malili lake system.

Fig. 4. Caridina dennerli from the Malili lake system. A. Distribution. B.-C. Colour pattern of living animals (not to scale). Pictures courtesy of Chris Lukhaup.

Fig. 63. Bayesian Inference phylogram (mtDNA, 16S and COI) of Caridina from the ancient lakes of Sulawesi (for other taxa compare von Rintelen et al., in review). Numbers on branches are, from top, Bayesian posterior probabilities, ML and MP bootstrap values. The scale bar indicates the number of substitutions per site. For each sequenced specimen museum accession numbers are provided. Locality abbreviations (compare Fig. 1): Lam (Lampesue River catchment), LT (Lake Lontoa), MHL (Lake Mahalona), MS (Lake Masapi), MT (Lake Matano), MTca (Lake Matano catchment), Pat (Patingko River), Pon (Ponsoa River), TW (Lake Towuti), TWca (Lake Towuti catchment), PS (Lake Poso), PSca (Lake Poso catchment), SSul (South Sulawesi), SESul (Southeast Sulawesi).

Fig. 64. Bayesian Inference phylogram (mtDNA, 16S and COI) showing taxa within the two Malili clades (for other taxa compare von Rintelen et al., 2009). For each sequenced specimen museum accession numbers are provided. Locality abbreviations (compare Fig. 1): Lam (Lampesue River catchment), LT (Lake Lontoa), MHL (Lake Mahalona), MS (Lake Masapi), MT (Lake Matano), MTca (Lake Matano catchment), Pat (Patingko River), Pon (Ponsoa River), TW (Lake Towuti), TWca (Lake Towuti catchment), SSul (South Sulawesi).