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Environ Biol Fish (2008) 81:7786 DOI 10.

1007/s10641-006-9173-6

ORIGINAL PAPER

Diel activity and reproductive territory of the Japanese bagrid catsh, Pseudobagrus ichikawai
Katsutoshi Watanabe

Received: 23 August 2005 / Accepted: 16 October 2006 / Published online: 22 November 2006 Springer Science+Business Media B.V. 2006

Abstract The endangered Japanese bagrid catsh, Pseudobagrus ichikawai, is typically nocturnal, emerging from shelters, such as crevices along the shore, at sunset to actively forage in open areas throughout the night with short-term movements of up to 40 m (except for mature males). Just before sunrise, individuals return to their shelters, from which they rarely emerge in daytime. Mature males (3 years-of-age or older) are found only in the vicinity of their shelters of during both day and night, which they defend with strong territorial behaviour. The restricted reproductive resources (shelters) probably severely limit the population size and reproduction in this species. Keywords Nocturnal Shelter Home range Siluriformes Bagridae

Introduction Diel activity patterns of shes, i.e., diurnal, nocturnal or crepuscular, are generally related to feeding modes and predator avoidance (Helfman 1978, 1993). Siluriforms (catshes) are predominantly nocturnal, likely depending on non-visual capabilities including tactile, acoustic, chemical or electrical sensory organs (Burgess 1989, Helfman 1993). Diel activity patterns of catshes have been quantitatively studied in the laboratory (Mashiko 1981; Boujardt 1995; Hossain et al. 1999) or based on temporal patterns of diel captures in the eld (e.g., Piet and Guruge 1997; Arrington and Winemiller 2003; Bhat 2003). The diel activity pattern or nocturnal habit of catshes in the eld, however, has rarely been investigated in detail (but see Vokoun and Rabeni 2006 using radiotracking). Bagrid catshes, comprising one of the largest families in the Siluriformes, are known to be generally nocturnal. In contrast to their nighttime activity, bagrid catshes hide in shelters such as crevices in riverbeds and banks (Kawanabe et al. 2001). Such shelters are also used as reproductive sites by some bagrid species (e.g., Nikolskii 1954). The quantity and utility of the shelters should affect population density and dynamics of the sh through reproductive success and carrying capacity. However, few observations have been conducted on the use of shelters by bagrids, as

K. Watanabe Laboratory of Ichthyology, Tokyo University of Fisheries, 4-5-7 Konan, Minato, Tokyo 108-8477, Japan Present Address: K. Watanabe (&) Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwakecho, Sakyo, Kyoto 606-8502, Japan e-mail: watanak@terra.zool.kyoto-u.ac.jp

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well as other catshes, especially in relation to diel changes. The Japanese bagrid, Pseudobagrus ichikawai, which is distributed in rivers within a restricted area in central Honshu and is now severely threatened, has been designated as endangered (category IB) in the Red Data Book of Japan (Watanabe 1997; Kobayakawa 2003). It has been threatened by human impacts, especially in the loss of shelters following the construction of concrete retaining walls in streams (Watanabe 1997, Kobayakawa 2003). Accordingly, detailed information on the spatio-temporal use of shelters and home ranges relative to diel activity patterns is necessary for evaluating the factors limiting the population size, and in ascertaining the extent of human impacts. This study was primarily aimed at clarifying the pattern of diel activity of P. ichikawai in a natural habitat. Furthermore, short-term movements and territorial behavior observed around male shelters were described in detail, those being important in relation to the ecological niche and conservation of the species.

vations were made around one of four patches of shelters inhabited by catsh, the patch being located in a 5-m section of the right bank protected by natural boulders (observation site by scan sampling for behavior, OS, see below; Fig. 1). Observations The survey was conducted from June 19 to July 29 in 1992, including the reproductive season of the catsh. The minimum and maximum water temperatures (WT) on each day were 1519C and 1926C, respectively. Sunrise was at 03:4004:10 and sunset, 19:4019:50 during the study period. The time difference (about 30 min) was not considered in the data analysis.

Materials and methods Study site The study was conducted in a tributary of the Nagara River system, Gifu Prefecture, central Honshu, Japan. Pseudobagrus ichikawai was distributed in the middle reaches (about 6 km) of this river. Observations were made in an 80-m slowwater reach (1750 m2), above a 1 m high weir, including a pool, slow run and glide (Fig. 1), where the density of catsh was relatively high (2.6 sh m-1). The maximum depth and current velocity were about 150 cm and less than 50 cms1 (mostly less than 10 cms1) in calm condition, respectively. The banks were comprised of bedrock, sand bars and articial protection using natural boulders or concrete, partly covered with overhanging or underwater vegetation. Migration of catsh between this 80-m reach and either the upper or lower reaches was very rare, according to a capture-recapture investigation (Watanabe unpublished data). Underwater obser-

Fig. 1 Characteristics of the study site in the Nagara River system, Gifu Prefecture, central Honshu, Japan. OS, site for behavior observations by scan sampling. Contour lines indicate water depth in cm

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The catsh were caught by hand net at the beginning, middle and end of the survey (each 23 days) and their standard length (SL) measured. The sh were sexed by the presence or absence of a male genital papilla posterior to the anus and a swollen abdomen in gravid females. The sh were divided into three age groups by sex (AS-groups: 1-year-of-age (+), 2+, 3+), based on size distribution (Fig. 2) as well as individual growth data obtained by a capture-recapture investigation conducted from 1990 (Watanabe unpublished data). The 2+ and older females and 3+ and older males had matured in 1992. The AS-groups could be easily identied during underwater observations. Over 80% of individuals (n = 137283, with 95% condence interval estimated from capture-recapture data; not shown) inhabiting the 80-m reach had been previously marked by a combination of injected colored liquid latex and a small piece cut from the adipose n, or by natural marks (e.g., scars on the body). Based on proportional changes in gravid females in the adult population (2+), spawning was

estimated as having started around July 12, peaking after July 18 and ending before July 27 (Fig. 3). The investigation included three sets of underwater observations. First, diel changes in activity, use of shelters and interactions between individuals were observed at the OS section (Fig. 4) by the scan sampling method (Martin and Bateson 1990) (termed as SCAN). Although the structure of crevice interstices at the OS was complicated and not recorded in detail, there seemed to be complex interconnections between the crevices, inferred from the tracking individual sh. The 5-m OS was divided into ve 1-m sections (IV), each being further divided into 4 (horizontal, AD) 2 (vertical) subsections (Fig. 4). Scans were done for each subsection in sequence over a 10-minute period (1070 scans) for each section. Standard length (SL), behavior (motionless, swimming, foraging or aggressive interactions) and individual identication (if possible) were recorded for each individual observed. Observations at nighttime were made using an unltered spotlight. When illuminated by a light, the catsh

Female
10 inds. 120 n = 60 n = 58 n = 80

Male
10 inds. n = 43 n = 38 n = 52 120

> 3+ Standard length (mm)


100 100

> 3+
80

2+

80

2+
60 60

Young (1+)
40 40

1921 June

23 July

27, 29

1921 June

23 July

27, 29

Fig. 2 Size distribution of Pseudobagrus ichikawai by sex during the study period. Age classes determined from distribution and growth data from marked individuals

Standard length (mm)

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Fig. 3 Rainfall and proportion of Pseudobagrus ichikawai females after spawning in during the study period

Fig. 4 Appearance of the OS section (upper) of the study site and relative density of Pseudobagrus ichikawai observed in each subsection (lower). AD, subsections. Blank,

<1.0%;*, 1.02.5%;**, 2.55.0%;***, >10.0%. b, booulder; c, cover; w; wood

5.010.0%;****,

usually remained in their positions and showed natural behavior, at least for a short time. They subsequently moved away from the light. Because the same sh was usually observed at the same position at the next scan, disturbance caused by the ashlight was considered to be negligible. SCAN observations were made for a total of 5530 min (107117 times for each section), on 24 days between June 22 and July 25, a total of 17,183 sh data being obtained. The frequency of agonistic interactions between individuals in each AS-group was corrected for visual counts of individuals in each group, by which counts of interactive actions were divided. Second, the diel patterns of locomotor or foraging activity in open areas were investigated by a visual census over the entire 80-m reach of the study area (CENSUS). The census was done along a regular course, following upstream (right side) and downstream (left side) directions, while counting the number of individuals in

each AS-group. Fifty census replicates were performed, taking 17 7 SD (range, 728) min/ census, between June 26 and July 10. Third, short-term movements of catsh were surveyed in the 80-m reach by mapping each individual sh with underwater visual observations and temporal capture of marked sh at night (PLOT). Eleven replicates of the PLOT investigation (each took 74 13 SD (range 4699) min) were performed between July 8 and 24. The location data of marked sh from the size distribution analysis and SCAN observations around the area of shelters were also included in PLOT data.

Results Diel activity During CENSUS, only ve or fewer P. ichikawai individuals were observed outside their shelters in

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daytime, whereas 50100 individuals were found at night-time, without distinct uctuations (oneway ANOVA, p = 0.137 among four 2-hourintervals between 20:00 and 3:00), showing a clear nocturnal activity pattern (Fig. 5). In daytime, most sh remained hidden in the interior part of the shelters, only 05 individuals (individuals)/m/
Fig. 5 Diel changes in the number of Pseudobagrus ichikawai individuals observed around the shelters by SCAN (plots) and in open areas by CENSUS (open circle with SD bar). h, section I; n, section II; + , section III; , section IV; , section V

scan being observed in SCAN (Fig. 5). In contrast, for a short time at sunset, the number of individuals observed around shelters dramatically increased to a maximum of 13 individuals/m/scan, sh swarming around the shelters and then gradually dispersing to open areas. At night-time, fewer than 2 individuals/m/scan were observed

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around the shelters. For a short time at sunrise, the catsh again gathered around the shelters, before retreating inside. On several occasions, just before sunrise at light levels darker than 0.1 lx, catsh were observed to rush back to the shelters simultaneously. A total of 69% sh were observed in sections III and IV during SCAN (Fig. 4). The diel activity pattern was especially distinct in younger sh (1+, 2+) (Fig. 5). Although older female sh showed a similar pattern, one or two individuals stayed at the entrance of their shelters even at night-time. In contrast, 3+ and older males, all of which had matured, did not show any obvious diel pattern, two or three individuals always swimming in and around the shelters. During daytime, all sh, except for 3+ males, shared the same shelters, irrespective of body size or sex, the frequency of attacks between sh being low (0.015 times/individuals/scan on average). On the other hand, the mature males (3+) showed an intense aggressive tendency (0.102 times/individuals/scan), defending a range of shelters against conspecic and other shes, as described below. Short-term movements During the PLOT investigation, a total of 45 individually identied sh were observed 5 or more times, and located at night and/or day (Fig. 6 for selected data). The data indicated that the short-term movements (home range) of the young and female catsh extended to 2040 m around the shelter (e.g., ID code, yc1L0L5, yc1L0L6). Some older females were found only in or near shelters, even at night (e.g., c4L0R4, c3L0L5), as mentioned above. Large males (3+ years or older) were all observed only around shelters during both day and night (Fig. 6). Aggressive interactions and reproductive territories No interactions between P. ichikawai individuals were recorded in the open areas at night, compared with frequent interactions, marked by aggressive behavior, observed around the shelters, especially when many sh occurred there around sunset and sunrise. A total of 456 aggressive interactions observed during SCAN included

four types of behavior: butting (79.8%), chase (12.7%), mutual circling (5.3%) and sideways body pushing (2.4%). Among these, mutual circling, observed mainly between two 2+ males with similar body size (42.8%), was the most violent behavior, in which each sh bit the caudal peduncle, pectoral n or basal bony part of the n of its opponent, or pushed heads, sometimes continuing for 20 or 30 s. Attacks directed to other sh species (cyprinids and a goby) represented 5.7% of all aggressive behavior. Of 456 aggressive interactions, 3+ and older males were the dominant attackers (57.3%; Table 1), especially during the last half of the observation period (after July 10: 69.6%), when reproductive activity was increasing (see Fig. 3). Such large males were swimming continuously around particular shelters, adopting very aggressive behavior against both conspecics and other shes in defense of their reproductive territories. Mainly butting or chase behavior occurred, when the large males detected other sh. The next most dominant attackers were 2+ males (20.1%). Most attacks were made by larger individuals against smaller ones, although the young sh (1+) were attacked infrequently (0.8%; Table 1). During the study period, seven individuallyidentied mature males (M1M7; 110123 mm SL, 3+ and older) were observed in the OS section, including one individual that maintained the same territory throughout the period (M4), one that changed the location of its territory (M3) and others that appeared only for short periods (M1, M7) (Fig. 7). The territorial males occupied shelters 12 m wide. A drastic change in the site occupied by each individual was observed between 10th and 13th of July, when spawning started (see Fig. 3). Intense aggressive interactions between mature males were also recorded at that time (Fig. 7).

Discussion Diel activity pattern and habitat use This study conrmed the typical nocturnal habit of P. ichikawai, individuals exhibiting all-night activity outside their shelters around the repro-

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Females ( > 3+)


Patch of shelters
c4L0R4 (10)

Day Night
20 m 0

Observed in shelter

Young (1+)
yc1L0L5 (11) yc1L0L6 (11) yc1L1L8 (19) yc1L2R5 (7)

Males (2+)
GL3L5 (7) GL6L7 (9) GL6R7 (8) GL6R9 (18)

c4GL2R9 (12)

GL0R8 (8)

GL1R6 (8)

GR1R3 (6)

m+ (>50)

Fig. 6 Sites where each marked individual were found. Only typical patterns are shown. Plots connected by a line indicate those on the same day. ID codes shown in italics with number of plots in parentheses

ductive season. Little change in the activity levels was apparent at night. The species is known to be a tactile (using barbels) benthivor-

ous feeder (Watanabe unpublished data), so individuals swimming out from their shelters at night could be searching for prey (and possibly

Table 1 Frequency (%) of aggressive interactions corrected by total number of individuals observed (n) Attacker n Opponent 1 1. Young (1+) 2. Females (2+) 3. Females (3+) 4. Males (2+) 5. Males ( 3+) 6. unknown Total 1235 5085 4453 3514 2078 818 17183 0.5 0.3 0.0 0.0 0.0 0.0 0.8 2 0.0 0.9 3.4 7.2 13.8 1.4 26.6 3 0.0 0.1 2.8 3.5 17.6 1.4 25.3 4 0.0 0.0 4.8 8.9 14.9 6.2 34.8 5 0.0 0.0 0.0 0.0 3.5 0.0 3.5 6 0.0 0.0 0.0 0.0 1.9 1.4 3.3 Other shes 0.0 0.0 0.0 0.0 5.7 0.0 5.7 0.5 1.3 11.0 19.7 57.3 10.3 100.0 Total

Females (2+)

Males ( >3+)
m(>50) m$ (21) PD-gr (>50)

c3L0L5 (30)

L2R0 (10)

L6L9 (6)

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Fig. 7 Positions of mature male individuals observed in the OS section between July 8 and 17. Each symbol (except star) indicates a different individual. See Fig. 4, for subsections. Large and small symbols in left panel indicate

observations 0.1 times/scan and <0.1 times/scan, respectively. Right panel shows a schematic diagram depicted from data in left panel. Star indicates aggressive interaction between mature males

mates by mature females). Casual observations have indicated that the nocturnal habit of the species occurred through the year except for winter when activity levels were very low (Watanabe 1994a). In the eld just before sunset, many individuals emerged from their shelters, and gradually swam away. They appeared hesitant to leave the shelters until conditions were completely dark. Just before sunrise while still dark (<0.1 lx), on the other hand, the catsh were observed returning to the shelters simultaneously. Experimental evidence has indicated that the activity of some catsh taxa is directly affected by light conditions, light inhibiting activity outside shelters (e.g., Mashiko 1979, 1981; Britz and Pienaar 1992; Hossain et al. 1999). Mashiko (1979) showed that

a Japanese bagrid, Pseudobagrus tokiensis (referred to as P. aurantiacus), a congener of P. ichikawai, was active without periodicity for an entire day under continuous darkness. This is probably also applicable to P. ichikawai. Mashiko (1979) indicated that the nocturnal activity of P. tokiensis terminated over a wide range of light intensity (104100 lx) in 1130C WT experimental conditions. Pseudobagrus ichikawai may also respond to very weak light conditions and quickly terminate activity before sunrise under natural conditions. Distinct ontogenetic changes or exibility in diel activity patterns were not observed in P. ichikawai, unlike other sh groups (e.g., Reebs 2004, Fraser et al. 2006). The typical nocturnal habit in P. ichikawai may represent a phyloge-

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netic constraint in siluriforms, which have developed several types of non-visual sensory organs, including barbels as tactile and chemical organs (Burgess 1989). However, from an ecological viewpoint, P. ichikawai should benet from nocturnal activity, because they can utilize benthic arthropods, many of which appear on the surface of river beds at night-time so as to avoid severe predation by abundant diurnal shes, such as cyprinids (see Allan 1995). On the other hand, predation pressure on P. ichikawai to alter its diel pattern may be limited. Potential night-time predators (Japanese eel, Anguilla janponica, and mitten crabs, Eriocheir japonicus, which occurred at low densities) are unlikely to be more important than potential daytime predators (several bird species, such as herons, family Ardeidae, and kingsher, Alcedo atthis). The maximum recorded density of P. ichikawai was 34 individuals/m (Watanabe and Ito 1999, Watanabe unpublished data), a generally much lower level than that of co-occurring dominant species, e.g., dark chub, Zacco temminckii and river goby, Rhinogobius umineus. However, P. ichikawai should not be considered a negligible component of local communities. Temporal partitioning of feeding activity can reduce potential interspecic competition in communities (Schoener 1974; Sale 1979). Nocturnal species often utilize aquatic insects active at nighttime, unlike diurnal shes. Fish composition in freshwater communities is known to affect the behavior and biomass of aquatic arthropods (e.g., Dahl and Greenberg 1996; Miyasaka and Nakano 1999) and also cause top-down effects on stream primary productivity (e.g., Huryn 1998; Nakano et al. 1999). The impact on temperate freshwater ecosystems by nocturnal shes such as P. ichikawai are not yet understood, but should be analyzed in the context of temporal resource partitioning and the community-level consequences (Helfman 1993). Reproductive territories of mature males Pseudobagrus ichikawai has been observed in aquariums to spawn in male shelters (Watanabe 1994b). Although direct observations of reproductive behavior was difcult in the eld because

of the concealed spawning sites (see Fig. 8), distinct territorial behavior around shelters by large males with dark nuptial body coloration was observed during this study. In contrast with the short-term movements (2040 m) of females and younger males, mature males were found only around their territories. Aggressive behavior between mature males was observed particularly just before the beginning of the spawning season, suggesting a shortage of sites for reproduction. Gaining and maintaining a reproductive territory are likely to be costly for males in such a situation. I found several dead adult males in the study site just after the reproductive season. The survival rate of 2+ and older P. ichikawai males is suggested to be lower than that of comparable females (Watanabe 1994a, unpublished data), possibly due to the severe aggressive activity, even before maturation in the males (2+), as observed in this study. A lack of potential reproductive sites, which must be only a part of the shelters used by the species, would limit the number of males participating in reproduction and cause severe competition for mates among males. Larger body size in males (see Fig. 2) appears to be a result of sexual selection via male-male competition. Also, the limitation of sites for reproduction may be one of the most important factors limiting population size in this species. Human impacts through the construction of concrete retaining wall often cause an extreme situation with loss of such sites, and are leading to population decline in this species.

Fig. 8 A male Pseudobagrus ichikawai, 93 mm SL, protecting its spawning site with eggs (enclosed by broken line) in section V (see text, Fig. 4), on July 7, 1997

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86 Acknowledgements I sincerely thank the late S. Watanabe and his family, and S. Ando for their help during the eld work, S. Mori, M. Nagoshi and Y. Taki for invaluable advice and discussion, and the Fisheries Union of the Upper Reaches of the Tsubo River, the Board of Education of Minokamo City and Gifu Prefecture, and the Japanese National Agency for Cultural Affairs for providing permission for the study.

Environ Biol Fish (2008) 81:7786 Japanred data book 2nd ednvol 4, Pisces blackish and fresh water shes. Japan Wildlife Research Center, Tokyo, pp 110111 (in Japanese) Martin P, Bateson P (1990) Measuring Behavior, an Introductory Guide. Cambridge Univ Press, Cambridge, p 238 Mashiko K (1979) The light intensity as a key factor controlling nocturnal action in the catsh, Pseudobagrus aurantiacus. Jpn J Ichthyol 25:251258 Mashiko K (1981) Periodic nocturnal activities in the catsh Silurus asotus in captivity. Jpn J Ichthyol 28:148156 Miyasaka H, Nakano S (1999) Effects of drift- and benthicforaging sh on the drift dispersal of three species of mayy nymphs in a Japanese stream. Oecologia 118:99106 Nakano S, Miyasaka J, Kuhara N (1999) Terrestrial aquatic linkages: riparian arthropod inputs alter trophic cascades in a stream food web. Ecology 80:2435 2441 Nikolskii GV (1954) Special Ichthyology. (English translation by J.I. Lengy & Z. Krauthamer. 1961. Israel Program for Scientic Translations, Jerusalem. 538pp.) Piet GJ, Guruge WAHP (1997) Diel variation in feeding and vertical distribution of ten co-occurring sh species: consequences for resource partitioning. Environ Biol Fishes 50:293307 Sale PF (1979) Habitat partitioning and competition in sh communities. In: Clepper H (ed) Predatorprey system in shery management. Sport shing institute, Washington, DC, pp 323331 Schoener TW (1974) Resource partitioning in ecological communities. Science 185:2739 Vokoun JC, Rabeni CF (2006) Summer diel activity and movement paths of athead catsh (Pylodictis olivaris) in two Missouri streams. Am Midl Nat 155:113 122 Watanabe K (1994a) Growth, maturity and population structure of the bagrid catsh, Pseudobagrus ichikawai, in the Tagiri River, Mie Prefecture, Japan. Jpn J Ichthyol 41:1522 Watanabe K (1994b) Mating behavior and larval development of Pseudobagrus ichikawai (Siluriformes: Bagridae). Jpn J Ichthyol 41:243251 Watanabe K (1997) Pseudobagrus ichikawai. In: Nagata Y, Hosoya K (eds) Circumstances in endangered Japanese freshwater shes and their protection. Midori Shobo, Tokyo (in Japanese), pp 122132 Watanabe K, Ito S (1999) Population size and distribution of Pseudobagrus ichikawai, an endangered bagrid catsh, in the Kawaura River. Jpn J Ichthyol 46:1530 (in Japanese with English abstract)

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