Corresponding author: Vladimir D. Ivanov (
Academic editor: Dominique Zimmermann
Caddisflies (
Caddisflies (order
The amazing species richness and endemism of the Malay Archipelago exists in one of the most geologically dynamic regions of our planet. The geological history of the Malay Archipelago is very complex (
A famous boundary dividing the Asian and Australian biotas, Wallace’s line, is one of the most disputed topics in biogeography since the end of the XIX century (
Malay Archipelago, sampling areas (rectangles), and the major biogeography divider lines:
The knowledge of the caddisflies of the Sunda Islands was poor for a long time. The first representative summary was given by Georg
Nevertheless, the caddisflies of the island of Lombok remained practically unknown, and this is still the state of knowledge of the caddisflies of the islands east of Lombok, including the Moluccas. This was the reason for the members of the Department of Entomology of the State University of St. Petersburg to make collections on the islands of Java, Bali, and Lombok in 2008 and 2009. The species which were new for science have been already described elsewhere (
Field collection included the well-known methods of light catch by small water-filled UV traps installed close to the water edge, net sweeping during daytime in the riparian vegetation, and hand picking. Only adults were collected; larvae and pupae were left for future investigations. The material is preserved in 70% ethanol and is stored in the collections of the Zoological Institute of the Russian Academy of Sciences in St. Petersburg. Some specimens are in the collection of the first author. Data from the faunistic publications and some material from the Malicky collection have been used for the analysis of zoogeography and species distribution.
Sampling localities were as follows:
LOCALITY 1: Lombok, Senaru, Sindanggala waterfall, Tiu Kelep waterfall and irrigation canal,
LOCALITY 2: Lombok, Sembalung Lawang,
LOCALITY 3: Lombok, vic. Kembangkuning, 7 March 2008,
LOCALITY 4: Lombok, vic. Kembangkuning, 2 km N Kotaraja,
LOCALITY 5: Lombok, Pelangan Barat, near mt. Embit,
LOCALITY 6: Bali, Gitgit waterfall,
LOCALITY 7: Bali, Munduk, Melanting waterfall,
LOCALITY 8: Java, Bogor, Sadame river, Botanical Garden,
LOCALITY 9: Java, Ciapus, Gunung Salak,
LOCALITY 10: Java, Cipanas, 30 km SE Bogor,
Recent collection sites cited as localities 1–10 in the text for Lombok (
List of species known from Java, Bali and Lombok, with new records
Abbreviations: B – Borneo, P – Peninsular Malaysia, S – Sumatra, Wd – wide distribution; m – males, f – females.
Taxa/family, species | Java | Bali | Lombok | other | Found in localities (Loc) |
---|---|---|---|---|---|
|
|||||
+ | + | Loc.1:5m,1f; Loc.7:1m | |||
+ | + | + | S, P | Loc.1:1m; Loc.3: (1f); Loc.6: 1m; Loc.7:1m | |
+ | |||||
|
|||||
+ | |||||
+ | |||||
+ | + | + | Loc.1:21m, 159f; Loc.3:2m, 5f; Loc.4:7m, 19f; Loc.6:17m, 26f; Loc.7:10f; Loc.8:3m; Loc.9:2m, 15f | ||
+ | |||||
+ | Loc.10:2m | ||||
+ | |||||
|
|||||
+ | |||||
+ | S | ||||
+ | |||||
+ | B, P, S | ||||
+ | Loc.4:12m | ||||
+ | Loc.7:1m, (5f) | ||||
+ | P, S | Loc.4:5m | |||
+ | S | ||||
+ | S | ||||
+ | P, S | ||||
+ | Loc.1:3m | ||||
+ | |||||
+ | Loc.1:1m | ||||
+ | |||||
+ | P, S | ||||
+ | B, P, S, Wd | ||||
+ | P, S | ||||
+ | |||||
+ | + | + | Loc.4:1m; Loc.8:16m, (numerous f); Loc.10:4m, (13f) | ||
+ | S, P, Wd | ||||
+ | Loc.4:2m | ||||
+ | + | S, P, Wd | |||
+ | P, S | ||||
+ | + | S | |||
+ | |||||
+ | B, P | Loc.1:4m; Loc.4:1m | |||
+ | Loc.4:1m, (1f) | ||||
+ | Loc.1:1m, (1f); Loc.4:1m | ||||
+ | + | S | Loc.1:17m, (3f); Loc.4:1m; Loc.10:1m, (2f) | ||
+ | |||||
+ | S | ||||
+ | |||||
+ | |||||
+ | + | ||||
+ | |||||
+ | + | + | S | Loc.1:2m; Loc.6:59m, numerous f | |
+ | + | + | Loc.1:1m | ||
+ | + | + | Loc.4:2m Loc.6:1m | ||
+ | |||||
+ | |||||
+ | Loc.4:1m | ||||
+ | + | ||||
|
|||||
+ | S | ||||
+ | |||||
+ | |||||
+ | + | + | Loc.1:4m, 41f; Loc.5:1m; Loc.6:15m, 68f; Loc.4:43m, 193f | ||
+ | |||||
+ | + | S | Loc.1:2m | ||
+ | P | ||||
+ | S | Loc.10:1m, (1f) | |||
+ | + | ||||
+ | + | ||||
+ | Loc.5:1m | ||||
+ | |||||
+ | + | P, S | |||
+ | Loc.7:1m, 2f | ||||
+ | |||||
+ | Loc.1:7m, 1f; Loc.3:1m | ||||
+ | |||||
+ | |||||
+ | |||||
+ | + | + | P, S, Wd | Loc.1:3m, 6f; Loc.3:2f; Loc.4:3f; Loc.7:5m, 3f | |
|
|||||
+ | |||||
+ | |||||
+ | S | ||||
+ | |||||
+ | Loc.9:1m | ||||
|
|||||
+ | + | Loc.10:1m | |||
+ | |||||
+ | S | ||||
+ | S, P | Loc.8:2m, (1f) | |||
+ | P, S | ||||
+ | + | S | |||
+ | S | ||||
+ | + | + | P, S, Wd | Loc.1:3m, 6f; Loc.3:2f; Loc.4:3f; Loc.7:5m, 3f | |
|
|||||
+ | + | + | S | Loc.1:27m, (23f); Loc.2:1m; Loc.3:9m, (1f); Loc.4:15m; Loc.7:9m, (4f); Loc.9:1m, (10f) | |
+ | + | + | Loc.5:3m; Loc.8:1m, (5f) | ||
+ | + | S, P | |||
+ | Loc.4:3m | ||||
+ | |||||
+ | P, S | ||||
+ | + | + | S | Loc.1:1m; Loc.4:4m; Loc.6:16m, 16f | |
+ | Loc.9:7m; Loc.10:5m, (3f) | ||||
+ | P | ||||
+ | P, S | ||||
+ | |||||
+ | + | Loc.7:6m; Loc.8:2m, (5f); Loc.9:3m, (9f); Loc.10:1m | |||
+ | |||||
+ | |||||
+ | + | Loc.1:16m; Loc.3:1m; Loc.6:1m | |||
+ | |||||
+ | + | Loc.1:47m, (9f); Loc.4:20m, (46f) | |||
+ | Loc.1:52m; Loc.2:1m | ||||
+ | Loc.1:1m; Loc.4:2m | ||||
+ | Loc.10:1m | ||||
+ | Loc.10:1m | ||||
+ | |||||
+ | Loc.7:5m, (3f) | ||||
+ | Loc.8:5m; Loc.10:3m | ||||
+ | Loc.9:5m, (7f) | ||||
+ | |||||
+ | |||||
+ | Loc.1:1m | ||||
+ | |||||
|
|||||
+ | + | P | Loc.6:11m | ||
|
|||||
+ | |||||
+ | + | + | P, S | Loc.1:5m, 2f | |
+ | K, S | ||||
+ | S | ||||
+ | B | ||||
+ | B, S, P, Wd | ||||
+ | |||||
+ | S | ||||
+ | + | B, S, P, Wd | |||
+ | P, S, Wd | ||||
+ | B, P, S, Wd | ||||
+ | B, S, | ||||
+ | + | P, S, Wd | Loc.7:2f | ||
+ | B, P, S | ||||
+ | P, S, Wd | ||||
+ | B, P, S | ||||
+ | |||||
+ | S | ||||
+ | |||||
+ | |||||
+ | |||||
+ | S | Loc.8:16m | |||
+ | |||||
+ | + | + | P, S | Loc.4:(5f); Loc.5:1m, 22f | |
+ | |||||
+ | + | S | Loc.6:(2f); Loc.7:8m, (40f) | ||
+ | B, S | ||||
+ | Loc.1:1m | ||||
+ | + | B, S, P | |||
+ | |||||
+ | + | ||||
+ | + | + | B, P, S, Wd | Loc.4:1m, (6f); Loc.5:7m, (25f); Loc.8:23m, (28f) | |
+ | Loc.1:2f | ||||
+ | Loc.1:8m, (227f); Loc.4:3m, (6f) | ||||
+ | + | Loc.6:1m; Loc.8:26m, (numerous f); Loc.9:1m, (12f); Loc.10:2m, (4f) | |||
+ | S | ||||
+ | Loc.1:8m, (20f); Loc.3:3m, (4f) | ||||
+ | P | ||||
+ | B? | ||||
+ | |||||
+ | |||||
+ | Loc.7:4m, (4f) | ||||
+ | + | S | Loc.1:1m; Loc.4:3m, (3f); Loc.6:1m | ||
+ | + | + | S, Wd | Loc.1:1m; Loc.4:2m; Loc.6:4m, (11f) | |
+ | Loc.1:6m, (3f); Loc.3:4m, (5f); Loc.4:1m | ||||
+ | Loc.3:1m | ||||
+ | |||||
|
|||||
+ | + | S | Loc.7:1m | ||
+ | B, S | ||||
+ | + | S | Loc.1:8m, 3f | ||
+ | S | ||||
+ | |||||
+ | Loc.1:1m, (1f) | ||||
+ | + | S | Loc.7:2m, 2f | ||
|
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+ | Loc.10:1m | ||||
|
|||||
+ | + | + | Loc.1:3m, 1f | ||
+ | + | S? | Loc.6:1m | ||
? | S | ||||
+ | + | + | S, P, Wd | Loc.4:1f; Loc.7:2m, 3f | |
|
|||||
+ | S, P | ||||
|
|||||
+ | + | S | Loc.1:14m, 9f; Loc.4:2m | ||
+ | + | S, P, Wd | Loc.4:4m; Loc.7:1m | ||
+ | S, P | Loc.1:3m, (5f) | |||
+ | S | ||||
+ | + | ||||
+ | S, P | ||||
+ | + | S | Loc.1:1m | ||
+ | + | + | Loc.4:1m; Loc.5:1m | ||
+ | + | S | Loc.4:2m, (4f); Loc.7:1m, (5f) | ||
+ | |||||
+ | S | ||||
+ | Loc.8:11m, (6f); Loc.9:2m, (5f) | ||||
+ | |||||
+ | + | Loc.4:2m, 9f; Loc.5:1m | |||
+ | + | Loc.3(1f); Loc.7:(16f) | |||
+ | S | ||||
+ | Wd | ||||
+ | + | + | B, P, S, Wd | Loc.4:1m, 6f | |
+ | + | B, S, P | Loc.4:1m | ||
+ | S | ||||
+ | B, P, S, Wd | ||||
+ | + | S, P, Wd | Loc.4:5f | ||
|
|||||
+ | B, S | ||||
+ | |||||
+ | |||||
+ | + | + | P, S | Loc.1:1f; Loc.4:1f; Loc.6:3m | |
|
|||||
+ | |||||
+ | S | ||||
Total: 202 species | 146 | 73 | 61 | Collected by V.Ivanov, S.Melnitsky, N. Kluge: 84 species |
Synopsis of the number of species in Thailand, Sumatra, Java, Bali, and Lombok (except
Taxa/family, genus | Thailand | Sumatra | Java | Bali | Lombok |
---|---|---|---|---|---|
|
|||||
|
37 | 7 | 3 | 2 | 2 |
|
|||||
|
3 | 2 | 2 | 0 | 0 |
|
14 | 5 | 4 | 1 | 1 |
|
|||||
|
33 | 14 | 5 | 3 | 2 |
|
2 | 0 | 0 | 2 | 2 |
|
1 | 1 | 1 | 0 | 0 |
|
4 | 4 | 1 | 1 | 0 |
|
5 | 1 | 1 | 0 | 0 |
|
32 | 11 | 5 | 3 | 2 |
|
2 | 0 | 1 | 0 | 0 |
|
5 | 3 | 1 | 0 | 4 |
|
13 | 5 | 2 | 2 | 0 |
|
9 | 1 | 2 | 3 | 1 |
|
2 | 1 | 1 | 1 | 1 |
|
7 | 0 | 4 | 2 | 2 |
|
|||||
|
60 | 21 | 10 | 8 | 3 |
|
9 | 3 | 3 | 0 | 1 |
|
|||||
|
8 | 1 | 0 | 0 | 0 |
|
|||||
|
13 | 14 | 2 | 2 | 0 |
|
20 | 6 | 1 | 0 | 0 |
|
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|
51 | 16 | 6 | 2 | 1 |
|
41 | 5 | 1 | 1 | 1 |
|
|||||
|
13 | 2 | 3 | 3 | 3 |
|
24 | 10 | 5 | 2 | 1 |
|
19 | 5 | 11 | 5 | 5 |
|
0 | 0 | 1 | 0 | 0 |
|
|||||
|
4 | 1 | 1 | 1 | 0 |
|
|||||
|
8 | 10 | 8 | 1 | 1 |
|
1 | 1 | 1 | 1 | 0 |
|
5 | 5 | 3 | 0 | 0 |
|
12 | 8 | 2 | 1 | 0 |
|
2 | 3 | 2 | 0 | 0 |
|
13 | 5 | 5 | 0 | 0 |
|
13 | 3 | 3 | 1 | 1 |
|
29 | 11 | 7 | 4 | 4 |
|
30 | 13 | 7 | 4 | 6 |
|
|||||
|
41 | 15 | 5 | 3 | 2 |
|
|||||
|
6 | 0 | 1 | 0 | 0 |
|
|||||
|
16 | 6 | 2 | 2 | 1 |
|
|||||
|
14 | 3 | 1 | 0 | 0 |
|
|||||
|
20 | 12 | 3 | 2 | 3 |
|
42 | 8 | 0 | 1 | 0 |
|
63 | 17 | 4 | 2 | 3 |
|
7 | 3 | 1 | 0 | 0 |
|
50 | 39 | 4 | 3 | 3 |
|
3 | 3 | 1 | 0 | 1 |
|
11 | 5 | 1 | 0 | 0 |
|
1 | 1 | 0 | 1 | 0 |
|
1 | 1 | 1 | 0 | 1 |
|
|||||
|
10 | 5 | 3 | 0 | 0 |
|
4 | 1 | 1 | 1 | 1 |
The caddisfly faunas of the three islands correspond quite well. It is evident from the
Surprises are also possible when species described from far distant regions will be found. For example,
Endemics of generic or even more-inclusive categories may scarcely exist on these islands. The genus
Continuous impoverishment of the Asiatic mainland faunas in the direction of the Lesser Sunda Islands is obvious (
Collecting intensity plays a major role in the knowledge of a fauna. Of Thailand and (northern) Sumatra it is well known, of Western Java and Bali it is relatively well known, too. For Lombok, the results of the first visit are being presented here. Our experience shows that the first intensive collections in an unknown area may yield about 75% unknown species, followed by the second visit results constituting about 20%, while later the percentage of the new species decreases dramatically. We conclude that each of Bali and Lombok may have less than 100 species in total. Such a dramatic decrease of the species numbers from the continent to the islands is not a result of poor collections. Instead it is caused by the island impoverishment and remote position from the continent. The number of caddisfly species in an area of similar size in Thailand is three times larger than in Sumatra. Supposed extreme faunal richness of rainforests does not apply to the aquatic insects, like caddisflies, of this region. To compare with extratropical Central and Southern Europe: about 300 (Austria, Switzerland, Germany, European Russia) to 400 species (Italy, Turkey) are known from these countries.
The well-known Wallace’s line which runs between Borneo and Bali on one side and Sulawesi and Lombok on the other side (
It is obvious that there is not an absolute border and the division is rather based on the deep sea waters between Bali and Lombok which remained deep even during the Pleistocene glaciations when the sea level was much lower (
Which
It is of highest zoogeographical interest to collect representative samples from the other Sunda Islands east of Lombok, and from the Moluccas. One may wonder that no such material is available in museums despite the intensive collecting activity of many other entomologists. The results of the faunal studies in Wallacea are also of high conservation importance. They can be used to steer both current and future efforts that try to preserve endemic flora and fauna of this uniquely diverse part of the biosphere.
The caddisfly fauna of the islands of Java, Bali and Lombok is largely homogenous, and there is no gap or border between Bali and Lombok which would be expected from the hypothesis of Wallace’s line. A continuous impoverishment of the caddisfly fauna from the Asiatic continent over the chain of islands is striking. This gradual impoverishment is observed in species number as well as in the numbers of genera or families. The
This research was supported by grants from the Russian Foundation for Basic Research, grants 11-04-00076 and the Federal Support Program for Leading Scientific Schools NSH-3332.2010.4. The authors are grateful to two reviewers for their valuable comments.