Interciencia Asociación Interciencia [email protected] ISSN (Versión impresa): 0378-1844 VENEZUELA

2000 Barry Chernoff / Antonio Machado Allison / Phil Willink / Jaime Sarmiento / Soraya Barrera / Naercio Menezes / Hernan Ortega OF THREE BOLIVIAN RIVERS: DIVERSITY, DISTRIBUTION AND CONSERVATION Interciencia, septiembre, año/vol. 25, número 006 Asociación Interciencia Caracas, Venezuela pp. 273-283

Red de Revistas Científicas de América Latina y el Caribe, España y Portugal

Universidad Autónoma del Estado de México FISHES OF THREE BOLIVIAN RIVERS: DIVERSITY, DISTRIBUTION AND CONSERVATION BARRY CHERNOFF, ANTONIO MACHADO-ALLISON, PHIL WILLINK, JAIME SARMIENTO, SORAYA BARRERA, NAERCIO MENEZES and HERNAN ORTEGA

he forested lowlands of eral ichthyological results, including a Fishes were collected us- most Amazonian tribu- breakdown by sub-basin; (ii) compare the ing a variety of nets and netting tech- taries remain poorly ex- diversity with those of other South Amer- niques. Each group was equipped with plored. Lists of freshwater fishes do not ican regions and discuss any distribu- seines (5m x 2m x 1.25cm, 5m x 2m x exist for more than one or two of the tional aspects of the data; (iii) discuss the .63cm, 1.3m x .7m x .37cm), dip nets and major arms of the Amazon, a feature that economic and conservation importance of experimental gill nets (40m x 2m, mon- was prominently absent from an other- the results (iv) consider threats to the ofilament, with five 8m panels, mesh size wise remarkable work on the fishes and richness; and (v) to propose rec- from 1.25cm to 6.25cm). Team 2 pulled an fisheries of the Rio Madeira (Goulding, ommendations for conservation. otter trawl (mouth 3m wide) with two 15kg 1981). In there have been at- doors where the depth of the water was tempts to bring together information for Methods >2m over sandy or muddy stretches in a Amazonian streams (e.g., Lauzanne et al., manner modified from that described by 1991). More recently, a well documented Two teams, three-four Lopez-Rojas et al. (1984). Additionally, fauna from the Rio Gaupore/Itenez was persons each, made collections between 4 one of the river pilots threw a 2m cast net provided by Sarmiento (1998). and 21 September, 1996. The teams were in some deeper lakes or cochas. Lauzanne et al. (1991) usually isolated and worked in different Fishes were preserved recorded 389 species of freshwater fishes tracts of the Upper Rio Tahuamanu for in buffered 10% formalin solution. All from the Bolivian Amazon to which Sar- the first five days. Eighty-five collection specimens captured at the same place and miento (1998) added an additional 21 stations were established, each receiving a time were maintained separately from all species bringing the total to 410 species. unique, sequential field number. The field other collected specimens. Larger speci- This work reports upon the ichthyologi- stations were enumerated separately for mens were tagged individually using fine cal results of the first AquaRAP carried each group, identified as P1 and P2. At wire and punched cardboard tags and ei- out in a relatively small area of the upper each station, longitude and latitudes were ther placed in large liquidpacks contain- Rio Orthon. Two teams of ichthyologists registered from hand held GPS units. ing formalin with other specimens or surveyed aquatic in the rivers At each field station a were skeletonized, soaked in 40% isopro- Tahuamanu and Manuripi and their tribu- number of ecological variables corre- panol and dried. All material was wrap- taries over 17 days in 1996. Amazingly, sponding to a description of the ped and shipped to Chicago for sorting, 313 species of fishes were recorded. This were recorded. These included the shore, identification and enumeration in the Di- value includes a spectacular number of substrate, type of habitat (e.g., river, lake, vision of Fishes, Department of Zoology, new records for Bolivia and several spe- flooded area, etc.) as well as the water Field Museum of Natural History cies new to science, including a new pi- type. The classification of water type (FMNH). Fifty percent of the specimens ranha, Serrasalmus. (black, white, turbid) was checked with are housed in the Museum of Natural The purpose of this con- the results obtained by the limnology History, La Paz, Bolivia; the remaining tribution is to: (i) make known the gen- group. specimens were shared among the partici-

KEY WORDS / Fhises / Diversity / Distribution / Conservation / Bolivia /

Barry Chernoff and Phil Willink: Field Museum of Natural History, 1400 Lake Shore Dr. Chicago, Illinois 60605-2496, USA. e- mail: [email protected] Antonio Machado-Allison: Laboratorio de Ictiología, Instituto de Zoología Tropical, Universidad Central de Venezuela, Caracas, Venezuela and Field Museum of Natural History. e-mail: [email protected] Jaime Sarmiento and Soraya Barrera: Museo Nacional de Historia Natural, La Paz, Bolivia. Naercio Menezes: Museu de Zoologia, Universidade de São Paulo, São Paulo, Brasil. Hernan Ortega: Museo de Historia Natural (UNMSM), Lima, Perú.

SEP 2000, VOL. 25 Nº 6 0378-1844/00/06/273-11 $ 3.00/0 273 pating institutions: FMNH; Museu Zoolo- gia do Universidade do São Paulo, Bra- zil; Museo de Historia Natural San Mar- cos, Perú and Museo de Biología de la Universidad Central de Venezuela. The identifications were made in a careful but relatively rapid fashion. General works such as Eigenmann and Myers (1929) or Gery (1977) were used but preference was always given to systematic revisions (e.g., Vari, 1992; Mago-Leccia, 1994) and recent species descriptions (Stewart, 1985) if available. In many cases speci- mens were compared to types or historic material referenced in the literature and housed at FMNH. However, some identi- fication to the level of species or even genus was not possible. To do so would represent a less than scholarly approach to the . Instead we rely upon Figure 1. General Map of the Rio Orthon Basin, showing the Collecting Stations on the Manuripi morphospecies – the number of distin- ( ) and Lower Tahuamanu (+) rivers (11º8-10'S to 67º33'W) guishable entities present in our samples. This bears the assumption that such The Rio Nareuda is a major tributary of well as in the Lower Tahuamanu, just discernable entities or morphospecies are the Rio Tahuamanu. above its mouth in the Rio Manuripi. River putative taxonomic entities (i.e., species). The gear effort used to conditions permitted trawling. We were careful to check for sexual and make the 85 field stations, were as fol- ontogenetic differences. All specimens lows: seine = 73; trawl = 6; gill net = 5; Results and Discussion were examined critically and identified to cast net = 2. This adds up to 86 because their lowest taxonomic level (Table I). one station P2-04 included both gill net Diversity and distribution: General Another issue inheres and seine collections. The gill nets were in the appropriate selection of taxa across usually set for several days. Because no The species richness of lists in order to judge new additional or striking differences were noted for day fishes was spectacular. A total of 313 new records. We chose a conservative and night samples within the gill nets, species were captured and identified. Be- approach and did not include all of the they were recorded as single stations. cause an additional 5% that could not be taxa that we had collected. We elimi- Due to low water con- identified unambiguously were discarded nated from comparison about 5% of ditions during the time of the field sam- from further evaluation, the actual num- those taxa whose identifications were am- pling, motors had only limited use in the ber of species in the entire region is even biguous or unknown in our list and in Upper Tahuamanu and could not be used greater. The fishes (Table I) included published lists. So for example, if Hemi- in the Rio Nareuda except for in the vi- members of all trophic or activity groups, grammus sp. or H. sp. 1 occurred in both cinity of its confluence with the Rio ornamentals (e.g., Abramites hypselono- lists, it was not counted as a similarity or Tahuamanu. During the first portion of the tus), food fishes (e.g., a difference because there is no way to expedition team 1 worked in the Upper fasciatum), as well as miniatures (Scol- ascertain that the taxonomic designations Tahuamanu and also in the lower end of oplax cf. dicra, <20 mm SL) and large represent the same biological entity. the Rio Muymanu whereas team 2 col- fishes (e.g., Prochilodus cf. nigricans, However, Gephyrocharax sp. occurs in lected in the Upper Nareuda as well as in Doras cf. carinatus, >200 mm SL). our list but only G. chapare is reported a number of small streams (garapés) that Together Lauzanne et by either Lauzanne et al. (1991) or Sar- drained independently into the Tahua- al. (1991) and Sarmiento (1998) record miento (1998). In this case we count the manu. The Upper Tahuamanu and the Up- 410 species from all rivers within the G. sp. as a new record because we com- per Nareuda systems as well as their Bolivian Amazon. The 313 species of pared our material to G. chapare and it tributaries were surrounded by terra firme. fishes, therefore, represents a fauna is different. The possible error in this lat- For the middle portion greater than 76% of the number of spe- ter case is identical to the possible errors of the expedition, both teams were cies previously reported from all other in a list containing misidentified species camped together but team 1 focused Amazonian tributaries within Bolivia. bearing specific epithets or not. upon the Lower Nareuda down to its The number of fishes discovered in a mouth in the Rio Tahuamanu. Team 2 relatively small section of the Tahuamanu Collecting Stations collected upstream and downstream from and Manuripi rivers is more than three this confluence to just below the village times that reported for the entire Beni- Eighty-five collections of Filadelfia; this region is referred to as Madre de Dios basin (n=101, Lauzanne were taken in the Tahuamanu and Ma- Middle Tahuamanu. Conditions in the et al., 1991), more than 1.3 times that re- nuripi river basins from the on the border Middle Tahuamanu were such that trawl- ported for the Río Guaporé/Itenez with Perú downstream to Puerto Rico ing was accomplished successfully. (n=246, Sarmiento et al., 1998) and al- (Figures 1 and 2). The entire region was For the last period both most equal to (96%) that found over the divided into the following five subre- teams camped on the Rio Manuripi up- entire Río Mamoré basin (n=327, Lau- gions based on habitat diversity: Upper, stream from Puerto Rico. Though both zanne et al., 1991). Furthermore, Santos Middle and Lower Tahuamanu, Upper groups worked independently, they covered et al. (1984) reported 300 species from and Lower Nareuda, and the Manuripi. the same territory in the Rio Manuripi as the lower Río Tocantins of ;

274 SEP 2000, VOL. 25 Nº 6 fact, it was representative of a more widespread fauna within Bolivia. The most obvious possibility is that we used trawls to sample the bottom communities for the first time within Bolivian fresh- waters. If the 91 novel records were largely due to trawl samples then the lists would not really be comparable. That is, we might expect to find similar bottom communities in other regions; the unique- ness of the Tahuamanu-Manuripi region would diminish even though its overall diversity would remain exceptional. How- ever, this scenario did not obtain. The trawls captured 53 species (17% of the total) but only 15 species were captured in trawls exclusively. Of these, 10 were new records for Bolivia (Table I). Thus, 81 of the 91 species newly reported for Bolivia were captured by traditional means used commonly in ichthyological sampling. We have no doubt that more careful sampling as exemplified in Figure 2. General Map of the Tahuamanu River Basin, showing the Collecting Stations on the the Guapore/Itenez by Sarmiento (1998) Upper Nereuda ( ), Lower Nereuda ( ), Upper Tahuamanu ( ) and Middle Tahuamanu ( ) (11º18- and in the Tahuamanu-Manuripi by the 25'S to 68º45'-69º02'W) AquaRAP team will lead to increasing the number of taxa found within Bolivia Stewart et al. (1987) reported 473 species documented fauna increases the total and will increase knowledge of the distri- in the Napo River Basin of ; number of species inhabiting the Bolivian bution patterns of the fishes. Nonetheless, Goulding et al. (1988) reported 450+ Amazon by ca. 22% to 501 and increases it seems unlikely that the unique charac- species from the Río Negro Basin of Bra- the total for all of Bolivia (using the pre- ter of the Tahuamanu-Manuripi fauna zil and Machado-Allison et al. (2000) vious total in Sarmiento, 1998) 16% to will loose its uniqueness. listed 136 species from the Río Cuyuní 641 species. In fact with continued basin in Venezuela. In each of these lat- The significance of the sampling, the ichthyofauna of the Tahua- ter cases, the areas surveyed exceed Tahuamanu and Manuripi rivers for the manu-Manuripi region will continue to vastly that of the Tahuamanu-Manuripi Bolivian ichthyofauna is now clear. This rise and the number of species new to region sampled in the rapid assessment. limited region contains 62.5% of all Bolivia will be expected to increase. We It is important to con- species known from the Bolivian Ama- base this assumption upon the species ac- sider that the number of species reported zon and 48.8% of all fishes known from cumulation curves (Figure 3). The rate of for both the Guapore/Itenez and Mamoré Bolivia. Furthermore, the 91 species accumulation of species new to the expe- drainages included headwater habitats found in the Tahuamanu and Manuripi dition had not diminished; the graph that usually contain fishes with restricted rivers that are not yet known elsewhere (Figure 3) displays no asymptote. During distributions. Even in the Upper Tahua- in Bolivia represent within-country “en- the last six days, even with both groups manu and Upper Nareuda, there were few demism” values of 18% and 14% relative working the same region of the Manuripi, habitats, perhaps the two garapés, that to the ichthyofaunas for the Bolivian we increased the known fauna by 63 spe- could correspond to headwater-like con- Amazon and Bolivia, respectively. cies. Species were being added at a rate ditions. Based upon the rela- of 10.5 species per day. Because of the paucity tively sparse information published for The total capture and of knowledge concerning the distributions the ichthyofauna of Bolivia, of regions accumulation rates were remarkably simi- of the freshwater fishes of South Ameri- with more than 50 species, only Lake lar for each of the two collecting teams ca, it is difficult to state with any cer- Titicaca has a higher within-country “en- (Figure 3). Each group captured more tainty the degree of endemism repre- demism” percentage. No other region than 200 species that were often comple- sented in samples from the upper Río within Bolivia is currently known to con- mentary. By the end of the sampling pe- Orthon. However, we have apparently tain as high a percent of the Amazonian riod, the species collected by the groups uncovered not only a region with excep- or total country fauna. The impressive differed as much as 50% which main- tional species richness but a region with values reported for the Río Guapore/ tained the steep slope of the total accu- an exceptional number of new records for Itenez basin within the Parque Nacional mulation curve. These data are evident in Bolivia (Table I). Using the conservative Noel Kempff reported upon by Sarmiento the differences of species captured by the approach discussed above, we document (1998) must now be adjusted downwards two teams when they were both sampling 91 species not previously recorded from due to the new species totals. The ad- in the Río Manuripi. The teams captured Bolivia. Of the 91, 45 species include justed values are well below those of the a total of 220 species in the Río Ma- taxa with some questions associated with Tahuamanu and Manuripi rivers. nuripi. Individually they netted 179 and their exact name as well as include a Potentially, the impres- 169 species based upon 17 and 18 col- number of new species (e.g., the new sive nature of the Tahuamanu-Manuripi lections, respectively. However, only 128 characid, Hysteronotus sp. 1). The newly ichthyofauna could be tempered if, in species were common to both sets of col-

SEP 2000, VOL. 25 Nº 6 275 lections, from which Simpson´s Index of Similarity is 75.7%. Thus, the additional effort increased the collected fauna by 62 species, or by 28.2% Analyses of these data argue strongly that continued collecting will increase the size of the fauna known from the Tahuamanu-Manuripi region. Because the species that represent new records for Bolivia comprise more than 29% of all species captured, we expect that continued collecting will continue to uncover new records for the country as well. These data continue to exemplify that the Upper Orthon Basin is extremely diverse and may prove more diverse than even the Rio Mamore basin, which has received vastly more collecting effort. The biogeographic rela- tionships of the Tahuamanu-Manuripi re- gion are difficult to ascertain, again be- cause of how little that we know in gen- eral. There does seem to be a mix of taxa representing three distinct distributional el- ements. The first comprises widespread amazonian lowland species from the north Figure 3. Species accumulation curves for fishes collected in the rios Tahuamanu, and the east of the Madeira basin: e.g., Nareuda, and Manuripi, Pando, Bolivia, 4-21 September, 1996. Serrasalmus rhombeus, Schizodon fascia- Symbols: Group 1 (squares), Group 2 (triangles), and combined (X). tum, Anodus elongatus, Microschemobry- con geisleri, Cetopsorhamdia fantasia, never reached the asymptote of the spe- Filadelfia and saw the following species Pimelodus cf. altipinnis, Tatia altae, Elec- cies accumulation curve, the number of for sale: Pygocentrus nattereri, Pseudo- trophorus electricus, and Rhabdolichops species represented in this region is pre- platystoma fasciatum, Prochilodus nigri- caviceps. The second incorporates species dictably larger than we can document at cans, Curimata spp., Hydrolycus pectora- found almost exclusively in black waters present. Furthermore, because the number lis, Plagioscion squamosissimus, Mylos- of the Guapore/Itenez system that derive of new records comprises 29% of the soma duriventre, and Myleus sp. We from the Brazilian Shield: e.g., Hypopygus fauna of this region, it is reasonable to were told by the fishermen that the fishes lepturus, Carnegiella strigata, Aphyo- expect new records as well. of highest commercial value were the ti- charax alburnus, Hemigrammus cf. unilin- All of this leads to the ger , Pseudoplatystoma, and the eatus, Pyrrhulina australe, Nannostomus conclusion that within Bolivia, the upper serrasalmines including both the piranhas trifasciatus, Potamorrhina latior, Tatia Rio Orthon basin must be considered as and the pacus. In our samples, we caught aulopygia, Corydoras hastatus, Cichla a potential hot spot for the species rich- many other species that are either con- monoculus, and Aequidens cf. tetramerus. ness of freshwater fishes. Conservation sumed for subsistence or caught for sales The third describes those species in the efforts are needed to preserve the unique according to our guides. That list in- small garapés that are most head-water character of this fish fauna. At a second- cluded the following: Anodus elongatus, like habitats: e.g., Hysteronotus sp., Bra- ary level, more field studies are needed Cichla cf. monoculus, Cochliodon cf. co- chychalcinus copei, Bryconamericus cf. to finish documenting the ichthyofauna chliodon, Crenicichla spp., Duopalatinus caucana, Creagrutus sp., Cyphocharax of this amazing region. sp., Hemisorubim platyrynchos, Hoplias spiluropsis, Piabucus melanostomus, Tyt- malabaricus, Hoplosternum thoracatum, tocharax tambopatensis, Corydoras tri- Economic importance Hypostomus spp., Leiarius marmoratus, lineatus, Imparfinis stictonotus, and Oto- Leporinus spp., Myleus sp., Pimelodus cinclus mariae. Many of the fishes spp., Pristobrycon sp., Rhamdia sp., The discussions above found in the Upper Rio Orthon basin are Schizodon fasciatum, Sorubim lima, and show the nature of the uniqueness of the valuable as commercial resources for Triportheus angulatus. In that list the ichthyofauna of the Tahuamanu-Manuripi food and for the ornamental fish industry. “spp.” refers to a number of species river basins. A relatively small region We present a short discussion in order to within the genus that are eaten. Interest- contains 62.5% of all the freshwater spe- stimulate immediate research both into ingly, the following armored are cies known from the Bolivian Amazon. the potential of these resources to pro- consumed in the Río Madeira basin of The fauna comprises 91 species that have vide an economic alternative to other ac- Brazil (Goulding, 1981) but were rejected never been recorded from Bolivia – a tivities that damage the environmental as a source of food by our fishermen: feature which does not seem to be an ar- character of the region. Doras cf. carinatus, Pseudodoras niger, tifact of sampling or collecting methods. During the AquaRAP and Liposarcus disjunctivus. The region seems to contain an assem- we employed local fishermen who were Our discussions with lo- blage of fishes that may uniquely com- subsistence fishermen throughout the year cal fishermen including others in Puerto bine widespread Amazonian species, with and who fished commercially during the Rico indicated that the commercial food blackwater Brazilian shield species in ad- seasonal migrations. We also witnessed fisheries are burgeoning in the Tahua- dition to head water species. Because we truck-side fish sales in Cobija and in manu and Manuripi rivers. It was not

276 SEP 2000, VOL. 25 Nº 6 possible to obtain from our fishermen or immediately both within the Upper Río main classes: i) flooded areas; ii) small from the truck-side sellers the annual to- Orthon as well as downstream toward the tributaries; and iii) main channels. The tals for weight or value of the catch. Río Madeira. Statements such as that by flooded areas comprise the most critical However, the notion of increasing annual Walters et al. (1982) advocating across and highly endangered areas, including landings would not seem to make sense the board increase in fisheries are prema- the varzea (flooded forest), cochas, relative to the local populations that we ture in advance of the data on native swamps, forest lakes, etc. These areas encountered and interviewed for two rea- stocks. provide nursery grounds for perhaps 66% sons: i) the population of Pando while Economic potential also of the species that we captured (Gould- increasing is doing so slowly; and ii) the exists for the establishment of a harvest- ing, 1980, 1981, Amazon; Lowe-McCon- Bolivian residents have largely settled the based ornamental fishery. The extreme nell, 1987 Amazon-Guianas; Machado- region from cattle rearing areas or from number of species, the number of cochas Allison, 1990, 1993 ). Further- La Paz and do not have strong traditions and inundated flooded habitats makes the more, many species, including the pacus, of fish consumption. Most of their de- Tahuamanu-Manuripi regions especially feed on the fruits and nuts, including the mand is for premium species, tiger cat- attractive. These habitats are easily col- Brazil nuts, dropped by the plants into fishes and pacus, species that are delicate lected while serving as natural critical the water (see Goulding, 1980, 1981, in flavor. Apparently, much of the catch rearing habitats for ornamental fishes. The Machado-Allison, 1990, 1993). Goulding is exported across the border to Brazil ornamental fishes ranged from the com- (1981) characterizes the Brazilian portion and this exportation appears to be un- mon (e.g., Moenhausia sanctofilamenae, of the Rio Madeira as having a relatively regulated. Sarmiento (1998) also noted a Hemigrammus ocellifer) to ornamentals narrow flood plane and margin. This is similar pattern of unregulated exportation that are more highly prized, including the certainly true for the rivers Tahuamanu, of several tons per month with a slightly folowing: Astronotus crassipinnis, Apisto- Muymanu, Nareuda and Manuripi in the increasing demand from populations liv- gramma spp., Aequidens spp., Satanoper- areas that we surveyed. Given such a ing along the Río Itenez. ca cf. acuticeps, Eigenmannia spp., Apter- narrow flood plain there is little buffer Peres and Terbourgh onotus albifrons, Gymnotus carapo, Hypo- between logging and ranching activities (1995), Goulding (1980, 1981) and Goul- pygus lepturus, Heptapterus longior, Im- and these critical flood zones. Extended ding et al. (1988) document not only the parfinis stictonotus, Microglanis sp., development from Puerto Rico, Filadelfia importance of rivers in structuring human Brachyramdia martae, Chirocerus eques, and Aceradero threaten this flood plain settlements throughout Amazonia but also Scoloplax cf. dicra, Peckoltia arenaria, zone. the increasing dependence of humans on Parotocinclus sp., Otocinclus mariae, Hy- In the Upper Tahua- aquatic resources for sustenance. At this postomus spp., Ancistrus spp., Agamyxis manu and Upper Nareuda there were time we cannot document the size of spe- pectinifrons, Acanthodoras cataphractus, many smaller streams (garapés) and cies specific harvests that are sustainable Brochis splendens, Corydoras spp., Buno- tributaries with both black and white wa- for the future. There is no accurate data cephalus spp., Dysichthys spp., Nannosto- ter conditions. These smaller habitats are on the nature of into the mus trifasciatus, Pyrrhulina spp., Carn- highly threatened by deforestation and Tauhuamanu-Manuripi region. Many of egiella spp., Tyttocharax spp., Poptella cattle ranching. A number of the garapés the commercially important species such compressa, Phenacogaster spp., Priono- crossing the main road and on cattle as the curimatids and prochilodontids brama filagera, Metynnis luna, Parago- ranches are completely denuded of ripar- move out of tributaries and forest habi- niates alburnus, Iguanodectes spilurus, ian vegetation. Furthermore, one team tats into the main rivers to at the Hemigrammus spp., spp. walked through dense forests into a num- beginning of the flooding cycle (Gould- Hysteronotus spp., Aphyocharax spp., and ber of forest streams and neither captured ing, 1981). While many of the larger cat- Leporinus spp. The most important areas nor saw any fishes, or fishes migrate upstream to spawn and ap- within the region surveyed for the orna- aquatic in crystal clear water with parently ascend the cataracts in the upper mental fishes are the Upper Río Nareuda sand bottoms. These streams were down- Madeira into Bolivian waters (Goulding, and the Río Manuripi (Machado-Allison et stream from a ranch that was recently 1981). The only relevant data that we al., 1999). In some cases, trapped interior cleared and burned. We could only collected was that the abundance of the flooded forest lakes can be harvested en- speculate that ashes which are toxic to most favored species was relatively low tirely because these ephemeral environ- aquatic organisms or pesticides poisoned (see Machado-Allison et al. 1999, on the ments either dry out completely or be- these streams. abundance and distribution). Even though come anoxic. However, in the more per- These upper areas are it was the dry season and at relatively manent habitats the reproductive and highly diverse. We captured 168 species low water, there were still many suitable population biologies of the ornamental in the Upper Tahuamanu and the Upper habitats for these species. The gill net fishes must be studied in order to support Nareuda; slightly more than half of all samples did not yield the quantity of in- a sustainable enterprise. the species we discovered. Seventy-one dividuals for the commercially important species were found to inhabit both the species that would indicate bountiful Critical habitats Upper Tahuamanu and Upper Nareuda populations, even though our fishermen but 30 species were found only in these guided us to their favorite areas (with We identified a number upper regions. These upper regions con- monetary exchange for the catch). In our of critical habitats that are required for tain habitats that are similar to headwater experience from similar habitats in other continued survival of freshwater fishes areas with a unique and diverse fauna. regions of the Amazon and Orinoco river and maintenance of the spectacular biodi- These are among the most threatened basins where fish populations are appar- versity. These are the same habitats that habitats due to logging, deforestation and ently healthy, the commercially important support the growth and reproduction of ranching. species are reasonably well represented in economically valuable species. The main river channels gill net samples even in the dry seasons. The habitats are de- are not habitats that are usually focused We only suggest caution. And we recom- scribed in detail in Machado-Allison et upon. However, as pointed out above, the mend that the stocks must be surveyed al., (1999). However, they fall into three principal channels also contain a diverse

SEP 2000, VOL. 25 Nº 6 277 fauna with a number of unique or rare el- flora and fauna and the habitats that sup- seen through fish communities. SPB Aca- ements (e.g., Cetopsorhamdia fantasia). port their survival and life histories. demic Publishing, The Hague, Netherlands. The Río Tahuamanu above Filadelfia Protection of aquatic re- 200pp. contains a number of rocky outcrops, in- sources is critical not only from the Lauzanne L, Loubens G, Le Guennec B (1991) cluding some small rapids (cachoeras) standpoint of conserving biodiversity but Liste commente des poissons de l’Amazonie bolivienne. Rev. Hydrobiol. Trop. 24: 61- and shallow regions with many downed also for protecting a critical resource for 76. logs. In the dry season the area near the people who live in the region. Many Lopez-Rojas H, Lundberg JG, Marsh E (1984) Aceradero was almost un-navigable be- of the fishes are currently used for sub- Design and operation of a small trawling cause of the tree trunks and rapids. Such sistence or commercial fishery and have apparatus for use with dugout canoes. North regions are prime candidates for spawn- a high value. The ornamental fishes also American Journal of Fisheries Management, ing areas of the commercially important, have a very high value. 4: 331-334. large, pimelodid catfishes (e.g., Pseudo- Lowe-McConnell RH (1987) Ecological Studies platystoma fasciatum, Leiarius marmora- ACKNOWLEDGEMENTS in Tropical Fish Communities. Cambridge tus, Barthem and Goulding, 1997). As University Press. development continues towards the Peru- The expedition that pro- Machado-Allison A (1990) Ecología de los vian border pressure may be exerted on duced the data reported upon was part of peces de las áreas inundables de los Llanos the river channelization to permit ship- the Aquatic Rapid Assessment Program de Venezuela. Interciencia 15(6): 411-423. ment of supplies or equipment up river. (AquaRAP) of Conservation International Machado-Allison A (1993) Los Peces de los and the Field Museum. We would like to Llanos de Venezuela: Un Ensayo sobre su Historia Natural. CDCH-UCV, Caracas. Conclusions and Recommendations express our gratitude, for their help in 143 pp. the organization, planning, and execution The region of the Río of the fieldwork, to Kim Awbrey, Adrian Machado-Allison A, Chernoff B, Willink P, Sar- miento J, Barrera S, Menezes N, Berth T Tahuamanu-Río Manuripi, upper Río Or- Forsythe, Lois “Lucho” James, Debra (1999) Diversity and Abundance of Fishes thon basin, Pando, Bolivia is a potential Moskovits, Guillermo Rioja, and Tatyana and Habitats in the Río Tahuamanu and Río hotspot for the diversity of freshwater Wachter. Leeanne Alonso and two Manuripi Basins (Bolivia). Acta Biol. fishes. A total of 313 species were dis- anonymous reviewers provided excellent Venez. 19(1): 17-50. covered in the region of which 91 spe- suggestions for improving the manuscript. Machado-Allison A, Chernoff B, Royero-León cies represented new records for Bolivia. We are grateful to Mary Anne Rogers R, Mago-Leccia F, Velásquez J, Lasso C, This region contains 62.5% of the fishes and Kevin Swagel for their help with López-Rojas H, Bonilla-Rivero A, Proven- zano F, Silvera C. (2000) Ictiofauna de la found in the Bolivian Amazon and processing and distributing the collections Cuenca del Río Cuyuní en Venezuela. In- 48.8% of the species found in the entire of fishes. Funding for fieldwork was gen- terciencia 25(1): 13-21. country. Based upon our analyses of the erously provided by the W. Alton Jones Mago-Leccia F (1994) Electric Fishes of the number of new records and the species Foundation and the Marshall Field Funds, Continental Waters of America. Biblioteca accumulation curve (Figure 3), we predict Department of Zoology, Field Museum. de la Academia de Ciencias Físicas, Mate- that the number of fish species within For their support of AquaRAP we thank máticas y Naturales, Volumen XXIX, Ca- this region will increase with continued the 3-M Corporation, J. Fahn, L. Lee, D. racas, Venezuela. 206 pp + Tables. sampling. and J. Shores, and Environmental and Peres CA, Terborgh JW (1995) Amazonian na- The region is further Conservation Programs and the Depart- ture reserves: an analysis of the defensibil- distinguished because the ichthyofauna ment of Zoology, the Field Museum. Par- ity status of existing conservation units and design criteria for the future. Conservation comprises a unique assemblage of spe- tial support of research at the Field Mu- Biology 9: 34-45. cies. There are three distinct biogeo- seum was also provided by a Bass Fel- Santos GM, Jegu M, Merona B (1984) Catalogo graphical elements as follows: i) wide- lowship to AMA. de peixes comerciales do baixo Tocantins. spread lowland Amazonian elements from Electronorte/CNPq/INPA, Manaus. 83pp. the north and east; ii) Brazilian Shield el- REFERENCES Sarmiento J (1998) Ichthyology of Parque ements from the Río Guapore/Itenez; and Nacional Noel Kempff Mercado. Pp. 168- iii) headwater elements. 180, Appendix 5. In: Killeen TJ, Schulen- Because this region is a Barthem R, Goulding M (1997) The Catfish berg TS (eds.) A biological assessment of potential hotspot for freshwater fishes we Connection : Ecology, Migration, and Con- Parque Nacional Noel Kempff Mercado. servation of Amazon Predators. Biology RAP Working Papers 10, Conservation In- are concerned that a plan be implemented and Resource Management in the Tropics ternational, Washington, D.C. to conserve this unique and valuable re- Series, Columbia University Press, New source. The region should be managed in York. Stewart DJ (1985) A new species of Cetop- sorhamdia (Pisces: ) from the a sustainable way that will protect the Eigenmann CH, Myers GS (1929) The Ameri- Río Napo Basin of Eastern Ecuador. Copeia biodiversity. We recommend that zones can . Memoirs of the Museum of 1985: 339-344. of critical habitats with varzea, cochas, Comparative Zoology, Harvard, 43 (Part 5): 429-574. Stewart DJ, Barriga R, Ibarra M. Ictiofauna de main channels and upland areas should la cuenca del Río Napo, Ecuador Oriental: be created that will protect the biodiver- Gery J (1977) Characoids of the World. T.F.H. lista anotada de especies. Revista del sity. Rather than the formation of a park, Publications, New Jersey. 672 pp. Politécnico 12: 9-63. authorities may want to consider multiple Goulding M (1980) The Fishes and the Forests: Vari RP (1992) Systematics of the Neotropical use zones with some habitats restricted Explorations in Amazonian Natural History. characiform genus Curimatella Eigenmann University of California Press, Los Angeles. and Eigenmann (Pisces: Ostariophysi) with for modification. Because the region has 280 pp. a narrow floodplain, special care should summary comments on the Curimatidae. Goulding M (1981) Man and Fisheries on an Smithsonian Contributions to Zoology, 533, be taken to minimize the impact on this Amazon Frontier. Dr. W Junk. The Hague, 48pp. important habitat. We recommend that 132 pp. Walters PR, Poulter RG, Coutts RR (1982) development of a management plan in- Goulding M, Carvalho ML, Ferreira EG (1988) Desarrollo Pesquero de la Región Amazónica clude educational programs concerning Rio Negro: rich life in poor water: Amazo- en Bolivia. London: Tropical Products Insti- the ecological relationships between the nian diversity and foodchain ecology as tute, Overseas Development Agency.

278 SEP 2000, VOL. 25 Nº 6 TABLE I FISHES FROM THE BOLIVIAN AMAZONIAN REGION (Explanation of localities and sources of information at end of table) TAXA Bolivia Noel Bolivian Madre Itenez TAXA Bolivia Noel Bolivian Madre Itenez AquaRAP Kempff Amazon de (or AquaRAP Kempff Amazon de (or Mercado Dios Guapore) Mercado Dios Guapore)

Rajiformes Bryconamericus cf. peruanus P - P -- Bryconamericus cf. peruanus ----- Potamotrygon cf. hystrix --P --Bryconamericus sp. P ---- Potamotrygon motoro P ----Bryconops cf. alburnoides --P - P Potamotrygon cf. motoro --PPP Bryconops cf. caudomaculatus - P --- Potamotrygon sp. - - P - P Bryconops melanurus - PP- P Potamotrygonidae sp. - - P - P Bryconops sp. - P --- Catoprion mento --P - P Lepidosireniformes Chalceus erythrurus -- - Lepidosirenidae PP Chalceus macrolepidotus --P - P Lepidosiren paradoxa - PP--Characidium bolivianum --P -- Clupeiformes Characidium cf. fasciatum - P --- Clupeidae Characidium sp. 1 P ---- Pellona castelnaeana --PPP Characidium sp. 2 P ---- Pellona flavipinnis --PPP Charax gibbosus PPP- P Cheirodon fugitiva ** ---- Engraulidae P Cheirodon piaba --P -- Anchoviella cf. carrikeri ** P ----Cheirodon sp. 1 P ---- Engraulidae sp. 1 - - P - P Cheirodon sp. 2 P ---- Engraulidae sp. 2 - - PP- Cheirodon spp. - P --- Cheirodon sp. - - P -- Anostomidae Cheirodon stenodon --PP- Abramites hypselonotus - --Cheirodontinae sp. P ---- P P Cheirodontinae sp. - --- Anostomus cf. gracilis --P - P P Anostomus cf. plicatus -- - Cheirodontinae sp. 1 - - P -- P P Cheirodontinae sp. 2 - -- Anostomus proximus --P - P P Anostomus taeniatus -- - Cheirodontinae sp. (gr. Aphyodite)- P --- P P Clupeacharax anchoveoides --- Laemolyta sp. ** P ---- PP Leporinus cf. cylindriformis - ---Colossoma macropomum --PPP P Creagrutus beni -- -- Leporinus fasciatus --P - P P Leporinus cf. fasciatus ----Creagrutus sp. 1 P ---- P Creagrutus sp. 2 ---- Leporinus friderici P - P - P P Leporinus cf. friderici - ---Creagrutus sp. 3 P ---- P Ctenobrycon spilurus Leporinus cf. nattereri ** P ---- PPPPP Leporinus pearsoni -- - Cynopotamus amazonus --P -- PP Cynopotamus gouldingi ** ---- Leporinus striatus --P -- P Leporinus trifasciatus -- - Engraulisoma taeniatum ** P ---- PP Eucynopotamus biserialis ** ---- Leporinus sp. nov. (amazonensis)- - P -- P Rhytiodus argenteofuscus -- Eucynopotamus sp. 1 - - P -- PPP Eucynopotamus sp. 2 - - - Rhytiodus lauzannei --P -- P P Rhytiodus microlepis -- - Galeocharax gulo P - PP- P P Gephyrocharax chapare -- -- Schizodon fasciatum PPPPP P Gephyrocharax sp. P ---- Characidae Gnathocharax steindachneri - P --- Acestrorhynchus altus --P - P Gymnocorymbus ternetzi --P - P Acestrorhynchus falcatus --PP- Gymnocorymbus thayeri --P -- Acestrorhynchus falcirostris --P - P Hemibrycon sp. - - P -- Acestrorhynchus guianensis --P --Hemigrammus cf. bellottii - P --- Acestrorhynchus heterolepis --P - P Hemigrammus lunatus PPPPP Acestrorhynchus microlepis --P - P Hemigrammus cf. marginatus --P -- Acestrorhynchus cf. minimus --P - P Hemigrammus cf. megaceps ** P ---- Acestrorhynchus sp. - P ---Hemigrammus ocellifer PP--- Aphiodite cf. grammica --PP- Hemigrammus cf. pretoensis ** P ---- Aphyocharax alburnus PPPP- Hemigrammus sp. - - P -- Aphyocharax dentatus P - P --Hemigrammus? sp. P ---- Aphyocharax paraguayensis --P --Hemigrammus cf. tridens - P --- Aphyocharax pusillus ** P ----Hemigrammus unilineatus --P -- Aphyocharax rathbuni - P ---Hemigrammus cf. unilineatus PP--- Aphyocheirodon sp. nov. - - P --Holobrycon pesu --P - P Astyanacinus cf. moori --P --Hyphessobrycon agulha ** P ---- Astyanacinus multidens --P --Hyphessobrycon cf. anisitsi ** P ---- abramis --P --Hyphessobrycon bentosi --P -- Astyanax cf. abramis P ----Hyphessobrycon cf. bentosi - P --- Astyanax bimaculatus --P - P Hyphessobrycon callistus - PP-- Astyanax cf. daguae --P - P Hyphessobrycon cf. gracilior ** P ---- Astyanax fasciatus --P --Hyphessobrycon cf. herbertaxelrodi - P --- Astyanax lineatus --P --Hyphessobrycon cf. heterorhabdus - P --- Astyanax cf. mucronatus --P --Hyphessobrycon cf. minimus - P --- Astyanax sp. P ----Hyphessobrycon cf. scholzei - P --- Astyanax sp. 1 - P ---Hyphessobrycon serpae --PP- Astyanax sp. 2 - P ---Hyphessobrycon? sp. P ---- Brachychalcinus copei P - P --Hyphessobrycon cf. tucunai PP--- cephalus --PPP Hysteronotus sp. 1 ** P ---- Brycon erythropterus --P --Hysteronotus sp. 2 ** P ---- Bryconacidnus ellisi --P --Iguanodectes spilurus PPP- P Bryconamericus bolivianus --P --Jobertina lateralis - P --- Bryconamericus cf. caucanus ** P ----Knodus breviceps - P -- Bryconamericus cf. pachacuti ** P ----Knodus cf. caquetae ** P ----

SEP 2000, VOL. 25 Nº 6 279 TABLE I (continued) FISHES FROM THE BOLIVIAN AMAZONIAN REGION (Explanation of localities and sources of information at end of table) TAXA Bolivia Noel Bolivian Madre Itenez TAXA Bolivia Noel Bolivian Madre Itenez AquaRAP Kempff Amazon de (or AquaRAP Kempff Amazon de (or Mercado Dios Guapore) Mercado Dios Guapore)

Knodus cf. gamma ** P ----Roeboides cf. myersii P ---- Knodus cf. heterestes ** P ----Roeboides sp. 1 P ---- Knodus cf. moenkhausii --P --Roeboides sp. 2 P ---- Knodus sp. P ----Roeboides sp. 3 P ---- Knodus sp. - P ---Roeboides sp. - - P - P Knodus sp. 1 - - P --Roestes molossus --P - P Knodus sp. 2 - - P - P affinis --P -- Knodus cf. victoriae ** P ----Salminus brasiliensis --P -- Markiana nigripinnis --P - P Serrasalminae sp. P ---- Megalamphodus megalopterus - P ---Serrasalmus compressus --PPP Megalamphodus sp. - P ---Serrasalmus eigenmanni --PPP Megalamphodus sp. - - PP- Serrasalmus elongatus --P - P Metynnis argenteus --P --Serrasalmus hollandi --P - P Metynnis hypsauchen --P - P Serrasalmus cf. hollandi P ---- Metynnis cf. hypsauchen --P - P Serrasalmus marginatus ** P ---- Metynnis cf. lippincotianus --P --Serrasalmus rhombeus P - PPP Metynnis luna ** P ----Serrasalmus sp. P ---- Metynnis cf. maculatus 1--P - P Serrasalmus sp. - P --- Metynnis cf. maculatus 2--P - P Serrasalmus spilopleura --PPP Microschemobrycon geisleri ** P ----Stethaprion crenatum PPP- P Microschemobrycon hasemani --PP- Tetragonopterinae sp. 1 P ---- cf. chrysargyrea ** P ----Tetragonopterinae sp. 2 P ---- Moenkhausia colletti PP---Tetragonopterus argenteus P - PPP Moenkhausia cf. colletti --P - P Tetragonopterus cf. chalceus --P - P Moenkhausia cf. comma ** P ----Thayeria boehlkei - PP- P Moenkhausia cf. cotinho --P - P Triportheus albus --PPP Moenkhausia dichroura PPPPPTriportheus angulatus PPPPP Moenkhausia grandisquamis --P - P Triportheus culter --P - P Moenkhausia jamesi --PPP Triportheus sp. P ---- Moenkhausia cf. jamesi P ----Tyttocharax madeirae PPP-- Moenkhausia cf. lepidura PP---Tyttocharax sp. nov. P ---- Moenkhausia cf. lepidura --P - P Tyttocharax tambopatensis ** P ---- Moenkhausia cf. megalops ** P ----Vesicatrus sp. nov. - - P -- Moenkhausia oligolepis - PP- P Xenurobrycon polyancistrus --P -- Moenkhausia sanctaefilomenae PPP-- Moenkhausia sp. 1 P ----Curimatidae Moenkhausia sp. 2 P ----Chilodus punctatus - PP- P Moenkhausia sp. 3 P ----Curimata roseni --P - P Moenkhausia sp. 4 P ----Curimata vittata --P - P Moenkhausia sp. 5 P ----Curimatella alburna P - P - P Moenkhausia sp. 6 P ----Curimatella dorsalis P - P - P Moenkhausia sp. 7 P ----Curimatella immaculata P - P - P Moenkhausia sp. 8 P ----Curimatella meyeri P - PPP Moenkhausia sp. - P ---Curimatopsis macrolepis - P --- Myleus sp. P ----Cyphocharax notatus - P --- Myleus cf. rubripinnis --P --Cyphocharax plumbeus --P - P Myleus tiete --P - P Cyphocharax cf. plumbeus P ---- Mylossoma aureum --P --Cyphocharax sp. P ---- Mylossoma duriventris P - PPP Cyphocharax sp. nov. - - P - P Odontostilbe cf. fugitiva --PP- Cyphocharax spilura - PPP Odontostilbe hasemani P - PP- Cyphocharax cf. spilura - P --- Odontostilbe piaba ** P ----Cyphocharax spiluropsis P ---- Odontostilbe paraguayensis ** P ----Cyphocharax cf. spiluropsis - P --- Odontostilbe sp. 1 P ----Eigenmannina melanopogon --PPP Odontostilbe sp. 2 P ----Potamorhina altamazonica P - PPP Odontostilbe sp. - - P --Potamorhina laitior P - PPP Parecbasis cyclolepis --PPP Psectrogaster curviventris P - P - P Paragoniates alburnus P - P --Psectrogaster essequibensis --P - P Phenacogaster cf. microstictus ** P ----Psectrogaster rutiloides PPP- P Phenacogaster cf. pectinatus ** P ----Steindachnerina bimaculata --P -- Phenacogaster sp. 1 P ----Steindachnerina binotata --P -- Phenacogaster sp. 2 P ----Steindachnerina dobula P - PP- Phenacogaster sp. 3 P ----Steindachnerina hypostoma --PP- Phenacogaster? sp. P ----Steindachnerina guentheri ** P ---- Phenacogaster sp. - P ---Steindachnerina leucisca ** P ---- Phenacogaster sp. - - P - P Steindachnerina sp. P ---- Piabucus melanostomus P - P - P Steindachnerina sp. - P --- Piaractus brachypomus --PP- Poptella compressa ** P ----Cynodontidae Poptella orbicularis - PPPPCynodon gibbus P - PPP Prionobrama filigera P - PP- Hydrolycus cf. armatus --PP- Pristobrycon sp. ** P ----Hydrolycus pectoralis ** P ---- Prodontocharax melanotus --P --Hydrolycus scomberoides --PPP Pseudocheirodon sp. - P ---Rhaphiodon vulpinus P - PP- Pygocentrus nattereri P - PPP Roeboides affinis --PPP Erythrinidae Roeboides cf. descalvadensis --P --Erythrinus erythrinus --P -- Roeboides myersii --PPP Erythrinus sp. - P ---

280 SEP 2000, VOL. 25 Nº 6 TABLE I (continued) FISHES FROM THE BOLIVIAN AMAZONIAN REGION (Explanation of localities and sources of information at end of table) TAXA Bolivia Noel Bolivian Madre Itenez TAXA Bolivia Noel Bolivian Madre Itenez AquaRAP Kempff Amazon de (or AquaRAP Kempff Amazon de (or Mercado Dios Guapore) Mercado Dios Guapore)

Hoplias malabaricus PPPPPTatia altae ** P ---- Hoplerythrinus unitaeniatus - PP--Tatia aulopygia PPP-- Tatia cf. intermedia - P --- Gasteropelecidae Tatia cf. perugiae ** P ---- Carnegiella marthae - P ---Tatia sp. - P --- Carnegiella myersi P - P --Trachelyopterus coriaceus --P -- Carnegiella schereri --P - P Trachelyopterus cf. galeatus P ---- Carnegiella strigata PP---Trachelyopterus cf. galeatus --P -- Gasteropelacus sternicla PPP--Trachelyopterus maculosus --P -- Thoracocharax securis --P -- Thoracocharax stellatus P - P --Callichthyidae Brochis britskii --P -- Hemiodontidae Brochis multiradiatus --P -- Anodus elongatus ** P ----Brochis splendens P - P -- Hemiodopsis cf. microlepis --P - P Callichthys callichthys PPP-- Hemiodopsis semitaeniatus --P - P Corydoras acutus P - P -- Hemiodopsis unimaculatus --PPP Corydoras aeneus PPP-- Parodon cf. carrikeri --P --Corydoras armatus --P -- Corydoras bolivianus --P -- Corydoras geryi --P -- Nannostomus sp. - P ---Corydoras hastatus PPP-- - P ---Corydoras cf. latus --P -- Nannostomus trifasciatus PP---Corydoras cf. loretoensis ** P ---- Nannostomus unifasciatus --P - P Corydoras cf. napoensis ** P ---- Pyrrhulina australe PPP- P Corydoras punctatus --P -- Pyrrhulina brevis - P ---Corydoras sp. P ---- Pyrrhulina vittata P - P - P Corydoras sp. - P --- Corydoras sp. 1 - - P - P Prochilodontidae Corydoras sp. 2 - - P -- Prochilodus nigricans --PPP Corydoras trilineatus ** P ---- Prochilodus cf. nigricans P ----Dianema longibarbis P - P - P Prochilodus sp. 1 - - P - P Hoplosternum littorale - PPP- Prochilodus sp. 2 - - P --Megalechis thoractus P - P -- Siluriformes Cetopsidae Ageneiosidae Cetopsis sp. - - P -- Ageneiosus brevifilis --PP- Hemicetopsis candiru --P -- Ageneiosus cf. caucanus ** P ----Pseudocetopsis plumbeus --P -- Ageneiosus dentatus --PP- Pseudocetopsis sp. P ---- Ageneiosus madeirensis --P - P Pseudocetopsis sp. - - P -- Ageneiosus sp. P ---- Ageneiosus sp. - - P --Doradidae Ageneiosus ucayalensis --P --Acanthodoras cataphractus ** P ---- Tympanopleura sp. P ----Acanthodoras spinosissimus - P --- Tympanopleura sp. - - P - P Agamyxis flavopictus --P -- Agamyxis pectinifrons ** P ---- Amblydoras hancockii - P --- Amaralia sp. - - P --Amblydoras cf. hancockii P ---- coracoideus ** P ----Anadoras cf. grypus ** P ---- Bunocephalus sp. 1 P ----Anadoras weddellii --P -- Bunocephalus sp. 2 P ----Astrodoras asterifrons P - P -- Bunocephalus sp. 3 P ----Doras cf. carinatus ** P ---- Bunocephalus sp. - P ---Doras eigenmanni P - P -- Bunocephalus sp. 1 - - P - P Doras fimbriatus --P -- Bunocephalus sp. 2 - - P --Doras punctatus --P - P Bunocephalus sp. 3 - - P --Doras sp. - - P -- Dysichthys bifidus ** P ----Hemidoras microstomus ** P ---- Dysichthys cf. aleuropsis ** P ----Megalodoras irwini --P -- Dysichthys cf. amazonicus ** P ----Opsodoras humeralis --P -- Dysichthys cf. depressus ** P ----Opsodoras cf. humeralis P ---- Xiliphius cf. melanopterus ** P ----Opsodoras stubelii --P -- Opsodoras cf. stubelii P ---- Astroblepidae Opsodoras sp. 1 - - P -- Astroblepus longiceps --P --Opsodoras sp. 2 - - P - P Astroblepus sp. - - P --Platydoras costatus P - P - P Pseudodoras niger P - PPP Auchenipteridae Pterodoras granulosus --PP- Auchenipterichthys thoracatus P - P - P Trachydoras atripes --P - P Auchenipterus nigripinnis --PPP Trachydoras cf. atripes P ---- Auchenipterus nuchalis --PPP Trachydoras paraguayensis P ---- Auchenipterus cf. nuchalis P ----Trachydoras cf. paraguayensis --P -- Centromochlus cf. heckelii ** P ---- Centromochlus sp. 1 - - P --Helogenidae Centromochlus sp. 2 - - P --Helogenes marmoratus - P --- Entomocorus benjamini P - P - P Epapterus dispilurus --P - P Hypophthalmidae Parauchenipterus striatulus --P --Hypophthalmus edentatus --PP- Pseudotatia? sp. - P ---Hypophthalmus marginatus --P --

SEP 2000, VOL. 25 Nº 6 281 TABLE I (continued) FISHES FROM THE BOLIVIAN AMAZONIAN REGION (Explanation of localities and sources of information at end of table) TAXA Bolivia Noel Bolivian Madre Itenez TAXA Bolivia Noel Bolivian Madre Itenez AquaRAP Kempff Amazon de (or AquaRAP Kempff Amazon de (or Mercado Dios Guapore) Mercado Dios Guapore)

Loricariidae Rineloricaria cf. lanceolata --P - P Ancistrus cf. bolivianus --P --Rineloricaria sp. P ---- Ancistrus cf. megalostomus --P --Rineloricaria sp. - P --- Ancistrus sp. 1 P ----Rineloricaria sp. - - P -- Ancistrus sp. 2 P ----Scoloplax cf. dicra ** P ---- Ancistrus sp. 3 P ----Scoloplax sp. - - P -- Ancistrus sp. 4 P ----Spatuloricaria cf. evansii --P -- Ancistrus sp. - P ---Sturisoma nigrirostrum P ---- Ancistrus sp. - - P --Sturisoma cf. nigrirostrum --P -- Ancistrus cf. temminckii --PPP Aphanotorulus frankei P - P --Pimelodidae Aphanotorulus unicolor ** P ----Brachyglanis? sp. ** P ---- Cochliodon cf. cochliodon ** P ----Brachyplatystoma filamentosum --P - P Cochliodon sp. 1 - - P --Brachyplatystoma flavicans --P -- Cochliodon sp. 2 - - P --Brachyrhamdia martae ** P ---- Crossoloricaria sp. ** P ----Callophysus macropterus --P - P Farlowella acestrichthys --P --Cetopsorhamdia phantasia ** P ---- Farlowella cf. oxyrryncha ** P ----Cheirocerus eques ** P ---- Farlowella sp. 1 P ----Duopalatinus cf. malarmo ** P ---- Farlowella sp. 2 P ----Hemisorubim platyrhynchos P - P - P Farlowella sp. - P ---Heptapterus longior ** P ---- Farlowella sp. 1 - - P --Heptapterus sp. P ---- Farlowella sp. 2 - - P --Imparfinis bolivianus --P -- Glyptoperichthys lituratus P - PPP Imparfinis cochabambae --P -- Glyptoperichthys punctatus --P --Imparfinis guttatus --P -- Hemiodontichthys acipenserinus P - P - P Imparfinis sp. P ---- Hypoptopoma joberti PPPPPImparfinis stictonotus P - P -- Hypoptopoma sp. ----Imparfinis cf. stictonotus - P --- P Leiarius marmoratus - -- Hypoptopoma thoracatum - PP- P P P Hypoptopomatinae sp. - ---Megalonema platanum --P -- P Megalonema sp. ---- Hypoptopomatinae sp. nov. ? - - P -- P Hypostomus bolivianus -- --Megalonema sp. nov. P ---- P Microglanis sp. ---- Hypostomus cf. chaparae --P -- P Hypostomus emarginatus -- - Microglanis sp. - - P -- P P Paulicea lutkeni -- - Hypostomus cf. popoi --PPP PP Hypostomus sp. 1 ----Phractocephalus hemioliopterus --P - P P Pimelodella cf. boliviana ** ---- Hypostomus sp. 2 P ---- P Hypostomus sp. 3 ----Pimelodella cf. chaparae --P -- P Pimelodella cristata - -- Hypostomus sp. 4 P ---- P P Hypostomus sp. - ---Pimelodella gracilis P - P -- P Pimelodella hasemani ** ---- Hypostomus sp. 2 - P --- P Hypostomus sp. 3 - ---Pimelodella cf. itapicuruensis ** P ---- P Pimelodella mucosa -- -- Hypostomus sp. 1 - - P -- P Hypostomus sp. 2 - - --Pimelodella roccae --P -- P Pimelodella serrata -- -- Hypostomus sp. 3 - - P -- P Pimelodella cf. serrata P ---- Hypostomus sp. 4 - - P - P Pimelodella sp. - - -- Hypostomus sp. 5 - - -- P P Pimelodidae sp. P ---- Lamontichthys filamentosus P ----Pimelodidae sp. - --- Lamontichthys cf. filamentosus -- -- P P Pimelodina flavipinnis --P -- Liposarcus disjunctivus P ----Pimelodus “altipinnis” ** ---- Liposarcus disjunctivus (sp. nov.) - - -- P P Pimelodus altissimus (sp. nov.) **P ---- Loricaria cf. simillima --P --Pimelodus armatus ** ---- Loricaria sp. ---- P P Pimelodus cf. blochii P ---- Loricaria sp. - P ---Pimelodus cf. maculatus-blochi -- Loricariidae sp. ---- PPP P Pimelodus ornatus --P -- Loricariinae sp. - P ---Pimelodus cf. pantherinus ** ---- Loricariichthys cf. maculatus -- P PPP Pimelodus sp. 1 P ---- Loricariichthys sp. P ----Pimelodus sp. 2 ---- Loricariichthys sp. - - P PPP Pimelodus sp. 3 P ---- Otocinclus mariae P - P --Pimelodus sp. 4 ---- Otocinclus cf. mariae - --- P P Pinirampus pirinampu --P - P Panaque sp. P ----Platysilurus barbatus -- -- Panaque sp. n 1 - - -- P P Pseudopimelodus zungaro --P -- Panaque sp. n 2 - - P --Pseudopimelodus sp. 1 - - P -- Parotocinclus sp. ** P ----Pseudopimelodus sp. 2 - - P -- Peckoltia arenaria P ----Pseudoplatystoma fasciatum PPPPP Peckoltia cf. arenaria --P --Pseudoplatystoma tigrinum --PPP Planiloricaria cryptodon P - P --Rhamdia quelen --P -- Pseudohemiodon cf. lamina ** P ----Rhamdia sp. P ---- Pseudohemiodon sp. 1 P ----Rhamdia sp. - P --- Pseudohemiodon sp. 2 P ----Rhamdia sp. - - P -- Pseudohemiodon sp. 3 P ----Sorubim lima P - PPP Pseudohemiodon sp. - - P --Sorubimichthys planiceps --P -- Pseudohemiodon thorectes --P -- Pterosturisoma microps --P -- Pterygoplichthys sp. - P ---Acanthopoma cf. bondi ** P ---- Rineloricaria beni - PP- P Apomatoceros sp. - - P -- Rineloricaria lanceolata PP---Gyrinurus batrachostoma --P --

282 SEP 2000, VOL. 25 Nº 6 TABLE I (continued) FISHES FROM THE BOLIVIAN AMAZONIAN REGION (Explanation of localities and sources of information at end of table) TAXA Bolivia Noel Bolivian Madre Itenez TAXA Bolivia Noel Bolivian Madre Itenez AquaRAP Kempff Amazon de (or AquaRAP Kempff Amazon de (or Mercado Dios Guapore) Mercado Dios Guapore)

Homodiaetus cf. maculatus --P --Beloniformes Homodiaetus sp. P ----Belonidae Homodiaetus sp. - - P --Potamorrhaphis cf. eigenmanni - P --- Ochmacanthus cf. alternus ** P ----Potamorrhaphis sp. P ---- Ochmacanthus sp. - P ---Potamorrhaphis sp. - - P - P Ochmacanthus sp. - - P - P Strongylura sp. - - P -- Paracanthopoma sp. - - -- P Synbranchiformes Plectrochilus sp. P ---- Plectrochilus sp. - - P --Synbranchidae Pseudostegophilus nemurus P - P --Synbranchus marmoratus PPP- P Trichomycteridae sp. - P ---Perciformes Trichomycterus cf. barbouri --P -- Trichomycterus sp. - ---Cichlidae P Aequidens dorsiger -- -- Tridentopsis pearsoni ** P ---- P Vandellia cirrhosa - --Aequidens sp. 1 P ---- P P Aequidens sp. 2 ---- Vandellia hasemani --P -- P Aequidens sp. 3 P ---- Gymnotiformes Aequidens cf. paraguayensis ** P ---- Apteronotidae Aequidens cf. tetramerus P ---- Adontosternarchus clarkae ** P ----Aequidens cf. tetramerus - P --- Adontosternarchus sachsi --P --Aequidens viridis - PP- P Apteronotus albifrons P - P --Aequidens cf. vittatus --P -- Apteronotus bonapartii P - P --Apistogramma inconspicua --P - P Apteronotus sp. - - P - P Apistogramma linkei P - P -- Porotergus cf. gimbeli --P --Apistogramma sp. 1 P ---- Porotergus cf. gymnotus --P --Apistogramma sp. 2 P ---- Sternarchorhynchus oxyrhynchus --P --Apistogramma sp. 3 P ---- Sternarchorhynchus sp. - - P --Apistogramma sp. 4 P ---- Apistogramma sp. 1 - --- Electrophoridae P Apistogramma sp. 2 - P --- Electrophorus electricus ** P ----Astronotus crassipinnis PPP- P Gymnotidae Batrachops sp. - - PPP Biotodoma cupido - PP- P Gymnotus anguillaris --P --Chaetobranchiopsis orbicularis - Gymnotus cf. anguillaris ---- P PPP P Chaetobranchus flavescens - PP- P Gymnotus carapo PPP- P Cichla monoculus - Gymnotus cf. coatesi ** ---- PPPP P Cichla cf. monoculus P ---- Hypopomidae Cichlasoma boliviense --P - P Cichlasoma severum ** ---- Brachyhypopomus brevirostris P ---- P Brachyhypopomus cf. brevirostris -- --Cichlidae sp. P ---- P Crenicara sp. - - -- Brachyhypopomus pinnicaudatus ** P ---- P Crenicara cf. unctulata ** ---- Brachyhypopomus sp. P ---- P Gymnotiforme - ---Crenicichla cf. heckeli ** P ---- P Crenicichla johanna -- - Hypopomus cf. artedi --P -- P P Crenicichla lepidota -- - Hypopomus sp. 1 - P --- P P Hypopomus sp. 2 - ---Crenicichla semicincta --P -- P Crenicichla sp. 1 ---- Hypopygus lepturus PP--- P Crenicichla sp. 2 P ---- Rhamphichthyidae Crenicichla sp. - P --- Gymnorhamphichthys hypostomus --P --Crenicichla sp. - - P - P Rhamphichthys rostratus - PPP- Geophagus megasema --P - P Rhamphichthys sp. - - P --Heros sp. - P --- Heros sp. - - Sternopygidae PPP Laetacara dorsigera - P --- Distocyclus conirostris P - P --Mesonauta festivus PPPPP Eigenmannia humboldtii P - P --Mesonauta cf. insignis ** P ---- Eigenmannia macrops ** P ----Microgeophagus altispinosa P - P - P Eigenmannia cf. trilineata ** P ----Satanoperca cf. acuticeps ** P ---- Eigenmannia virescens PPP- P Satanoperca jurupari --PP- Rhabdolichops caviceps ** P ----Satanoperca pappatera --P - P Rhabdolichops troscheli --P --Satanoperca sp. P ---- Sternopygus macrurus PPPPP Eleotridae Cyprinodontiformes Eleotridae sp. - - -- Rivulidae P Cynolebias sp. - P ---Sciaenidae Pterolebias sp. - - P --Pachypops sp. - - P -- Rivulidae sp. - P ---Pachyurus sp. P ---- Rivulus sp. P ----Plagioscion squamosissimus P - PPP Rivulus sp. - P --- Rivulus sp. - - --Pleuronectiformes P Soleidae Achirus achirus - PP--

P: Presence; -: Absence of given species; **: Species collected by AquaRAP and believed to be new records for Bolivian Amazon. Locality information for fishes in Noel Kempff Mercado is from Sarmiento (1998). Locality information for fishes in the Bolivian Amazon, Madre de Dios, and Itenez (or Guapore) columns is taken from Lauzanne et al. (1991). Note that Madre de Dios and Itenez (or Guapore) are subsections of the Bolivian Amazon, and that the Bolivian Amazon includes additional basins not indicated in this table.

SEP 2000, VOL. 25 Nº 6 283