Caridina loehae 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 : 377-381

Caridina loehae Woltereck, 1937a

( Figs. 18-19 , Table 8)

Caridina Loéhae Woltereck, 1937a: 222 , Figs. I.5 a-d, pls. 3,6 (type locality: Matanno [Matano] islands, Lake Towuti XX, Lake Towuti at Loéha Island and at Lingkona).

Caridina loéhae – Woltereck, 1937b: 304, fig. 9; Fernandez- Leborans et al., 2006b: 1985, Table I (partly as C. loeha , erroneous spelling).

Caridina loehae – Chace, 1997: 13; von Rintelen et al., 2008: 2244, Table 1; Cai et al., 2009: 27, Fig. 7 (type locality of neotype: Lake Towuti, about 3 km south of Timampu, estuary of Sungei [River] Batuopa).

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

Material examined. – Lake Matano : 8 ex. ( ZMB 29062, n=8), north shore, 02°26.36'S, 121°19.03'E, loc. 84-03, on rocks, coll .

K. & T. von Rintelen, 1 Oct.2003; 20 ex. ( MZB Cru 1768, n=10; ZMB 29066, n=10, some SEM material), north shore, 02°27.28'S, 121°21.21'E, loc. 98-03, on rocks, coll. K . & T. von Rintelen, 5 Oct.2003; 3 ex. ( ZMB 29079, and few juveniles), south shore, Soroako , Salonsa , INCO boat house, 02°30.71'S, 121°20.45'E, loc. 04-05, on rocks, coll. K . & T. von Rintelen, 3 Jan.2005; 42 ex. ( MZB Cru 1769, n=30, ZMB 29080, n=12, some SEM material), south shore, Soroako, Salonsa , INCO boat house, 02°30.71'S, 121°20.45'E, loc. 19-03, on rocks, coll. K . & T. von Rintelen, 19 Sep.2003; 15 ex. ( MZB Cru 1770, n=7; ZMB 29118, n=8, some SEM material), south shore, east of Soroako, just west of Cape Patipuoho, 02°31.57'S, 121°23.41'E, loc. 99-03, on rocks, coll. K . & T. von Rintelen, 5 Oct.2003; 4 ex. ( ZMB 29233, some SEM material), southwest shore, Cape Nikomene , 02°32.24'S, 121°24.76'E, loc. 46- 03, on rocks, coll. K . & T. von Rintelen, 18 Sep.2003; 3 ex. ( ZMB 29244), north shore, 02°29.75'S, 121°25.81'E, loc. 42-03, on rocks, coll. K . & T. von Rintelen, 17 Sep.2003; 10 ex. ( MZB Cru 1771, n=5; ZMB 29168, n=5), north shore, 02°26.274'S, 121°18.83'E, loc. 133-04, on rocks, coll. K . & T. von Rintelen, 22 Jul.2004; 15 ex. ( MZB Cru 1772, n=7; ZMB 29223, n=6), north shore, 02°27.311'S, 121°21.047'E, loc. 09-05, on rocks in shallow water, coll. K . & T. von Rintelen, 7 Jan.2005; 2 ex. ( ZMB 29241), north shore, 02°25.67'S, 121°16.54'E, loc. 65-03, on mixed substrate, coll. K . & T. von Rintelen, 25 Sep.2003; 1 ex. ( ZMB 29447), south shore, 02°27.84'S, 121°13.88'E, loc. 63-03, on rocks, coll. K . & T. von Rintelen, 24 Sep.2003; 1 ex. ( MZB Cru 1773), south shore, at small islands, 02°28.476'S, 121°15.64'E, loc. 138-04, on rocks, coll. K . & T. von Rintelen, 24 Jul.2004; 2 ex. ( MZB Cru 1774), north shore, 02°27.418'S, 121°21.533'E, loc. 10-05, on mixed substrate, coll. K . & T. von Rintelen, 7 Jan.2005; 1 ex. ( MZB Cru 1775), south shore, canal between island and mainland, 02°28.46'S, 121°15.83'E, loc. 62-03, on rocks, coll. K . & T. von Rintelen, 1 Oct.2003; 1 cl (mm) 2.2-2.9 2.6 ± 0.2 2.6 27 rl / cl 0.5-1.3 0.8 ± 0.1 0.8 29 n dorsal rostral teeth 14-20 17 ± 2 17 29 n ventral rostral teeth 1-8 4 ± 2 4 29 abds6 / cl 0.5-0.8 0.6 ± 0.0 0.7 27 abds6 / abds5 1.5-2.1 1.8 ± 0.2 1.9 20 abds6 / h tel 0.9-1.7 1.0 ± 0.2 1.0 20 h tel / w tel 3.1-3.5 3.3 ± 0.2 3.3 6 n spines uropodal diaeresis 9-14 12 ± 2 11 5 h ch1 / w ch1 1.9-2.7 2.1 ± 0.2 2.1 27 h ch1 / h ca1 0.9-1.8 1.1 ± 0.2 1.0 30 h ca1 / w ca1 2.5-4.7 3.2 ± 0.5 3.3 26 h ch2 / w ch2 2.3-3.1 2.7 ± 0.2 2.6 26 h ch2 / h ca2 0.6-1.0 0.7 ± 0.1 0.6 30 h ca2 / w ca2 5.4-8.1 6.7 ± 0.7 6.7 27 n spines p3 2-6 3 ± 2 3 5 n spines p5 12-16 15 ± 2 16 5

ex. ( MZB Cru 1776), south shore, 02°27.85'S, 121°13.87'E, loc. 125-04, on rocks, coll. P. Koller & K. von Rintelen, 1 Aug.2004; 36 ex. ( MZB Cru 1780, n=14; ZMB 29084, n=22) , Petea River , 02°32.64'S, 121°29.51'E, loc. 101-03, on rocks, coll. K. & T. von Rintelen, 6 Oct.2003; 27 ex. ( MZB Cru 1781, n=13; ZMB 29460, n=14) . 02°32.672'S, 121°30.137'E, loc. F4-04, substrate unknown, coll. F. Herder, 25 Mar.2004.

Lake Towuti – 1 ex. ( MZB Cru 17771), outlet bay, at Cape Kombe, 02°48.083'S, 121°23.049'E, loc. 118-04, on mixed substrate, coll K. & T. von Rintelen, 29 Jul.2004 ; 8 ex. ( MZB Cru 1779), Larona River , close to outlet bay, 02°45.06'S, 121°20.12'E, loc. 50-03, on rocks, coll. K. & T. von Rintelen, 21 Sep.2003 .

Description. – Carapace length 2.2-2.9 mm (n=27). Rostrum ( Fig. 19A,B ; Table 8) short, reaching near or to end of second segment of antennular peduncle, 0.5-1.3 times as long as carapace (n=29), armed dorsally with 14-20 teeth (including 3-5 teeth posterior to orbital margin), armed ventrally with 1-8 teeth. Antennal spine situated below inferior orbital angle. Pterygostomial angle broadly rounded. Eyes well developed, anterior end 0.5-0.7 times length of basal segment of antennular peduncle (n=5). Antennular peduncle 0.8-1.1 times as long as carapace (n=5), second segment 1.6-1.8 times length of third segment, third segment 0.3-0.4 times length of basal segment. Stylocerite reaching 0.9-1.0 times length of basal segment of antennular peduncle (n=5). Scaphocerite ( Fig. 19D ) 3.7-4.7 times as long as wide (n=5).

Sixth abdominal somite 0.5-0.8 times length of carapace (n=27), 1.5-2.1 times as long as fifth somite (n=20), 0.9-1.7 times length of telson (n=20). Telson ( Fig. 19F,K ) 3.1-3.5 times as long as wide (n=6), distal margin rounded, without projection, with 3-4 pairs of spinules and 1 pair of dorsolateral spinules; distal end with 3-4 pairs of spines, lateral pair distinctly longer than intermediate pairs. Preanal carina ( Fig. 19C ) rounded, without a spine. Uropodal diaeresis ( Fig. 19E ) with 9-14 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 maxilliped 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. 19 N-P); chela of first pereiopod 1.9-2.7 times as long as wide (n=27), 0.9-1.8 times length of carpus (n=30); tips of fingers rounded, without hooks; dactylus 1.4-1.6 times as long as palm (n=6); carpus 2.5-4.7 times as long as wide (n=26), 1.2-1.3 times length of merus (n=5). Chela of second pereiopod 2.3-3.1 times as long as wide (n=26), 0.6-1.0 times length of carpus (n=30); tips of fingers rounded, without hooks, dactylus 1.6-2.1 times as long as palm (n=6); carpus 5.4-8.1 times as long as wide (n=27), 1.3-1.5 times as long as merus (n=6).

Third pereiopod ( Fig. 19G,H ) slender, dactylus 2.7-3.6 times as long as wide (terminal spine included, without spines of flexor margin; n=5), terminating in one large claw with 2-6 accessory spines on flexor margin; propodus 10.2-17.2 times as long as wide, 3.7-7.3 times as long as dactylus; carpus 4.6-6.1 times as long as wide, 0.6-0.7 times as long as propodus, 0.5-0.7 times as long as merus; merus 6.8-9.6 times as long as wide, bearing 1-4 strong, movable spines on posterior margin of outer surface.

Fifth pereiopod slender ( Fig. 19I,J ), dactylus 3.2-4.0 times as long as wide (terminal spine included, without spines of flexor margin; n=5), terminating in one large claw with 12- 16 accessory spines on flexor margin; propodus 12.8-20.3 times as long aswide, 5.2-8.1 times as long as dactylus; carpus 5.8-7.8 times as long as wide, 0.5-0.6 times as long as propodus, 0.5-0.7 times as long as merus; merus 7.9-12.6 times as long as wide, bearing 2-3 strong, movable spines on posterior margin of outer surface.

Endopod of male first pleopod ( Fig. 19L ) elongated triangular, 1.8-2.3 times as long as proximally wide (n=5), without appendix interna. Appendix interna of male second pleopod ( Fig. 19M ) 0.8-0.9 times length of appendix masculina (n=5).

Ovigerous females with 10- 19 eggs (n= 4 females); egg size 0.8-1.1 x 0.5-0.7 mm (n=60, eggs with and without eyes).

Distribution. – C. loehae is endemic to the Malili lake system. There, however, widely distributed in Lake Matano and in Petea River, but was only found in the outlet bay area of Lake Towuti ( Fig. 18A ). Woltereck mentioned the occurrence of C. loehae at three other localities in Lake Towuti, e.g. at Loeha Island, therefore the distribution shown here may not be complete.

Biology and ecology. – C. loehae is a typical hard substrate dweller, mainly occurring on and under smaller rocks, and on gravel in shallow water (above 5 m), where it is often collected with other rock dwellers such as C. parvula and C. dennerli . When disturbed, it tries to escape side- or downwards and often stays attached to rocks. The red colour can turn blue in stressed animals.

Colour pattern. – Body colouration (including appendages) of C. loehae varies from light to dark red with three conspicuous white transversal stripes in regular intervals on the posterior part of the carapace and the abdomen, a scattering of small white spots covering the whole body and uropods with white tips ( Fig. 18B ), although the white colour is not always strongly pronounced. Eggs were observed to be red.

Taxonomic remarks. – Cai et al. (2009: 19) stated: “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”. Consequently, they designated a neotype for C. loehae from Lake Towuti (ovigerous female, cl 2.6 mm, ZRC) (2009: 27) .

With regard to its small size (carapace length 2.2-2.9 mm, median 2.6 mm), C. loehae is one of the smallest species in the Malili lake system, similar to C. spongicola and C. parvula . It differs from both by the generally more slender and fragile rostrum, a different rostrum length (reaching near or to end of second segment of antennular peduncle vs. longer in C. spongicola ), and a lower number of spines on the dactylus of the fifth pereiopod (12-16, median 16 vs. 21-31, median 27 in C. spongicola and 34-39, median 37 in C. parvula ). With regard to the rostrum, it also resembles C. masapi . However, C. loehae is generally smaller (vs. 2.1-4.6 mm, median 3.1 mm in C. masapi ), has a shorter rostrum (not overreaching end of second segment of antennular peduncle vs. longer in C. masapi ), and a lower number of spines on the dactylus of the fifth pereiopod (vs. 30-44, median 33 in C. masapi ).

In the molecular phylogeny ( Figs. 63-64 ), C. loehae is genetically distinct from all other ancient lake species.

Figures

Fig. 18. Caridina loehae from the Malili lake system. A. Distribution. B. Colour pattern of living animals (not to scale). Pictures courtesy of Chris Lukhaup.

Fig. 19. Caridina loehae from the Malili lake system. A. Cephalothorax and cephalic appendages, female (ZMB 29066); B. Woltereck’s drawings of the rostrum, from two lakes (modified from 1937a); C. Preanal carina; D. Scaphocerite; E. Uropodal diaeresis, F. Telson, female (ZMB 29080); G. Dactylus of third pereiopod, female (ZMB 29118); H. Third pereiopod; I. Dactylus of fifth pereiopod; J. Fifth pereiopod; K. Distal end of telson, female (ZMB 29080); L. Endopod of male first pleopod (ZMB 29066); M. Appendix masculina of male second pleopod; N. SEM image of chela and carpus of first and second pereiopods, female (ZMB 29233); O. First pereiopod; P. Second pereiopod. Scale bars: A, D = 1.0 mm; C, F, H, J, L-P = 0.5 mm; E, G, I, K = 0.1 mm; B = no scale available.

Fig. 5. Caridina dennerli from the Malili lake system. A. Cephalothorax and cephalic appendages, female (ZMB 29049); B. Uropodal diaeresis, male (ZMB 29051); C. Telson, male (ZMB 29156); D. Scaphocerite, male (ZMB 29051); E. Preanal carina, F. Third pereiopod, female (ZMB 29326); G. Fifth pereiopod; H. Distal end of telson male (ZMB 29156); I. Dactylus of third pereiopod female (ZMB 29326); J. Dactylus of fifth pereiopod; K. Endopod of male first pleopod (ZMB 29051); L. Appendix masculina of male second pleopod; M. First pereiopod, male (ZMB 29156); N. Second pereiopod; O. SEM image of anterior part of chela, male (ZMB 29104); P. SEM image of chela and carpus of first and second pereiopods, female (ZMB 29024). Scale bars: A, D = 1.0 mm; C, E-G, K-P = 0.5 mm; B, H-J = 0.1 mm.

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. 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).