Galaxea, Journal of Coral Reef Studies (Special Issue): 243-252（2013）

Abstract The Banggai cardinalfish Pterapogon kauderni (Koumans, 1933) is a reef-associated fish endemic to the Banggai Archipelago in Central Sulawesi. Considered en­dangered due to exploitation for the marine aquarium trade, introduced populations have become established along trade routes, including in Palu Bay. A national action plan has been developed to ensure P. kauderni conservation with a sustainable use approach. Over the period 2007-2009 research aiming to support the plan was conducted into the biology and ecology of both endemic and introduced P. kauderni populations. Aspects covered include micro­habitat use; feeding habits; reproduction; the effects of salinity on juvenile P. kauderni growth and survival rates. Ontogenetic shift in micro­habitat use was observed in both endemic and introduced populations. Sea anemones, threatened in both the en­demic and introduced habitat, may play an important role in recruit survival. The presence of juvenile urchins (Diadema sp.) in P. kauderni stomach contents indicates that the relationship may be more complex than was previously thought and an ontogenetic shift in diet is indicated. Unlike other Apogonidae, observed P. kauderni gonads were single lobed. Female gonad development indicated a peak in reproductive activity around the full moon continuing into the third quarter. Males may mature slightly earlier than females and observations made raised questions relating to sex ratio and sex determination. Ju­

venile P. kauderni growth was highest at 27 ppt salinity, with a significant reduction in long term survival rates at salinities below 24 ppt. The results have implications for the sustainable management of Banggai cardinalfish pop-ulations and habitat.

Keywords Banggai, micro­habitat, diet, reproduction, sa­linity

Introduction

The Banggai cardinalfish Pterapogon kauderni (Koumans, 1933)  is  a  small  fish  (around  6 cm  maximum  standard length  SL)  of  the  family Apogonidae,  first  collected  by Walter Kaudern (1917-1920) and subsequently described by Frederick Koumans as belonging to a new Genus. Pre­viously unexploited, collection for the marine ornamental trade began in the 1980's and expanded rapidly after the “rediscovery” of P. kauderni by Gerald Allen in 1994. Introduced populations have been established along trade routes due to intentional and accidental releases. The en­demic distribution according to Vagelli (2008) and known introduced populations (Erdman and Vagelli 2001, Ndobe et al. 2005, Moore and Ndobe 2007a, Lilley 2008) are shown in Fig. 1. Found in shallow coastal waters to a

The Banggai cardinalfish: an overview of local research (2007-2009)

Samliok NDOBE1, Abigail MOORE2, *, NASMIA1, MADINAWATI1, Novalina SERDIATI1

1 Aquaculture Study Program, Tadulako University, Kampus Bumi Tadulako, Palu 94118, Central Sulawesi, Indonesia2 Sekolah Tinggi Perikanan dan Kelautan (STPL), Kampus Madani, Jl Soekarno­Hatta, Palu 94118, Central Sulawesi, Indonesia

* Corresponding author: A. MooreE­mail: abigail2105@yahoo.com

Proc 2nd APCRS

Ndobe et al.: The Banggai cardinalfish: an overview of local research244

depth of around 5 m, including coral reefs, reef flats and seagrass beds, P. kauderni is a paternal mouthbrooder with direct development, predominantly diurnal and car­nivorous (Vagelli and Erdman 2002). Considered endan­gered by overexploi­tation, in 2007 P. kauderni was pro­posed for listing in CITES (Convention on the Interna­tional Trade in Endangered Species of Wild Flora and Fauna) Appendix II (CITES 2007) and was subsequently listed as Endangered in the IUCN (International Union for the Conservation of Nature) Red List (Allen and Donald­son 2007). The CITES proposal was withdrawn. Indonesia made a commitment to ensure sustain­able management of the species and a national multi­year multi­stakeholder Banggai Cardinalfish (BCF) Action Plan was developed (Ndobe and Moore 2009). Building on research begun in 2004 (Ndobe and Moore 2007), this paper presents an overview of some of the research carried out to support the development of conservation/sustainable use of P. kauderni, in line with the BCF Action Plan and forth­coming national legislation.

Materials and Methods

Research locationsField research was carried out at Tinakin Laut, a site in

the Banggai Archipelago (endemic distribution, around S 1°36′  and E 123°29′)  and at Mamboro  in Palu Bay  (in-troduced population, around S 0°47′53″ and E 119°52′). Both locations can be seen in Fig. 1. Ex situ research took place in the Laboratory facilities of Tadulako University (UNTAD) and the Sekolah Tinggi Perikanan dan Kelautan (STPL),  a private marine and fisheries higher education institute, both in Palu, the capital of Central Sulawesi Province.

Micro-habitat useQuantitative observations were made in 2007, using

belt (20×5 m) transects, following methods previously used (e.g. in Ndobe et al. 2005) with four habitats, four replicates and 4 micro­habitat categories. Three habitat types  were  defined  at  Tinakin  Laut,  all  with  abundant Diadema urchins: coral reef (CR), rubble dominated reef flat (RF) and seagrass (SG) while at Mamboro (MB) the habitat was  uniform:  reef  flat with  corals  and  seagrass, very few Diadema urchins. The micro­habitat categories were: Diadema sea urchins (D); sea anemones (A) ; hard coral (HC); and other (OT), comprising all other micro­habitat (e.g. seagrass, soft corals). P. kauderni associated with each micro­habitat within each transect were counted using 3 size categories based on Standard Length (SL): Recruits (SL＜15 mm); Juveniles (15 mm ≤ SL ≤ 35 mm)

Fig. 1　P. kauderni known endemic distribution (dotted line) and introduced populations (stars)

Ndobe et al.: The Banggai cardinalfish: an overview of local research 245

and Adults (SL .＞35 mm). In addition, qualitative obser­vations were made from 2007 to 2009 using a swim sur­vey method (Hill and Wilkinson 2004).

Feeding habitsAdult and sub­adult P. kauderni from Mamboro (30

specimens captured at random) were immediately sacri­ficed and weighed. The abdominal  cavity was dissected and guts were observed to estimate gut fullness, then they were removed and weighed. The Gut Fullness Index (GFI) used is shown in the legend of Fig. 3. The contents of each gut were placed in a test tube with a solution of 70% alcohol and gently shaken to separate the components. The components were then selected using a pipette and placed on microscope slides. A binocular microscope with a magnification factor of 10×4 was used to examine each item. Identification was based mainly on Sachlan (1972) supported by a number of other references.

ReproductionAdult and sub­adult specimens (SL 29-66 mm) were 

captured at random from the Palu Bay population at four phases of the moon: dark/new moon (day 29-1 of the lunar cycle), half­full waxing moon (day 7-8), full moon (day 14-16)  and  half-full  waning  moon  (day  22-23). Twenty specimens were taken at each phase, N＝4×20＝80.  Each  specimen was  sacrificed, weighed  then  dis-sected and the gonad observed to determine the Gonad Maturity Index (GMI) based on Effendie (2002) with a qualitative scale from I to IV as follows: I. Unripe. Gonad very small, sometimes semi­transparent; II. Beginning to mature. Gonad filling around a quarter of the body cavity; III. Nearly ripe. Gonad filling around half the body cavity; IV. Ripe. Gonad filling around three quarters of the body cavity, white fluid exuding from testes, eggs clearly visi-ble in ovaries. The gonad was then removed and weighed to determine the Gonadosomatic Index (GSI)＝gonad weight/total weight. For specimens with immature gonads for which the sex (ovary or testes) was not readily apparent with the naked eye (GMI I and in some cases GMI II), sex was determined through histological examination. A thin layer of gonad tissue was prepared and stained following the protocol described in Zairin (2004) and observed under a binocular microscope with a magnification factor 

of 10x4.

The effects of salinityJuvenile P. kauderni (15-18 mm SL) from the Palu Bay

population were reared for 60 days in aquaria at a density of 5 fish/10litres (0.5/l) provided with aeration and shelter. The research  took place  in  two stages.  In 2008, five sa-linity treatments were used (27 ppt, 29 ppt, 31 ppt, 33 ppt and 35 ppt), each with 4 replicates., while in 2009, six sa­linity treatments were used (16 ppt, 18 ppt, 20 ppt, 22 ppt, 24 ppt  and  26 ppt),  each with  3  replicates. The  juvenile fish were fed ad libitum with Artemia salina 3 times a day. Length and weight were measured at 10 day intervals and treatments/replicates were compared using the analysis of variance (ANOVA) and Tukey Test methods as described in Gasperz (2001).

Results

Micro-habitat useMicro­habitat use data are shown in Fig. 2. The age/size

class composition of P. kauderni at Tinakin Laut was similar for all three habitats, with a lower proportion of adult fish compared  to  the Mamboro population  (“% of sample”  data  set  in  Fig.  2)  and  all  fish  observed  were associated with one of the three main micro­habitat types. Diadema density was low at the Mamboro site (5-6/100  m2 as opposed to over 150/100 m2 at Tinakin Laut and all P. kauderni habitat surveyed in the Banggai Archipelago) and “other” micro­habitat consisted mainly of soft corals and seagrass. All recruit groups comprising three or more individuals as well as many juvenile P. kauderni were found in sea anemones, sheltering between the tentacles, often  co-habiting  with  clown  fish  (Amphiprion sp). No adults and large juveniles associated with sea anemone habitat were observed in close contact with or among the tentacles of the sea anemones.

During swim surveys, adult P. kauderni (SL＞3.5 cm) were occasionally observed close to anemones, including solitary brooding males with active embryos in their mouth pouches. No adults and large juveniles associated with sea anemone habitat were observed in close contact with or among the tentacles of the sea anemones. All size

Ndobe et al.: The Banggai cardinalfish: an overview of local research246

classes were observed associated with Diadema sp., but the largest recruit group observed in Diadema micro­habitat  consisted  of  3  individuals. Adult  fish  and  large juveniles were frequently associated with hard corals, while only 3 recruits (recorded at the Mamboro site) have been observed to date in this micro­habitat.Significant habitat and micro-habitat degradation were 

observed  or  reported,  including  destructive  fishing  and coral mining, predation by Acanthaster plancii, and over­fishing including a sharp increase in collection of anem-ones and Diadema urchins. Reduced P. kauderni popu­lation densities and very low numbers of recruits were observed after sea anemones had been harvested by local communities for human consumption (Tinakin Laut) and for ornamental use (Mamboro).

Feeding habitsGut content was dominated by zooplankton. Gut full­

ness index (GFI) data is shown in Fig. 3 and gut compo­sition data for adult and sub­adult P. kauderni is shown in Fig. 4. No empty or nearly empty guts (GFI 0 or 5) were

observed. The two brooding males in the sample had GFI values of 20 and 40. The diet of sub-adult fish was domi-nated by Crustacea (comprising over 80% of recognizable material) with an average of 3 components per gut. The diet  of  adult  fish  tended  to  be  more  varied.  Juvenile Diadema urchins and phytoplankton were found in both sub­adult and adult guts.

ReproductionGonad development data related to size (based on

standard length, SL) are shown in Table 1 and lunar peri­odicity data for female P. kauderni are shown in Fig. 5. Female gonad maturity peaked during full moon into the third quarter/waning and was lowest during the dark/new moon phase (GSI and GMI). There was no clear pattern in male gonad maturity. A single gonad was apparent in all specimens  (Fig.  6),  the  female  gonad  being  generally darker in colour. In female P. kauderni with GMI of III or IV, an egg mass was generally observed (19-102 eggs), separate from the gonad and encased in a white egg en­velope, with an overall appearance and containing an average number of eggs  (56) very  similar  to  that of  the egg masses observed in the mouths of brooding males.

There were more males than females in the (randomly obtained) sample for each moon phase (average sex ratio M:F＝1.6:1). The smallest male (SL＝29 mm) observed with GMI ≥ II was smaller than the smallest female (SL ＝33 mm). The smallest male (SL＝43 mm) with GMI III was larger than the smallest female with GMI III (SL＝ 41 mm) and smaller than the smallest female with fully

Fig. 2　Ontogenetic composition (percent) of P. kauderni observed in each micro­habitat

Fig. 3　Gut Fullness Index (GFI) of P. kauderni speci­mens examined

Ndobe et al.: The Banggai cardinalfish: an overview of local research 247

ripe gonad GMIV (SL＝45 mm).

The effects of salinityGrowth and survival rate data are shown in Fig. 7. The

highest growth rate for juvenile P. kauderni was obtained at 27 ppt salinity, with 100% survival rate. Survival rate was high (90-100%)  for  all  treatments  for  the  first  10 days. Over a 60 day period, below 24 ppt the reduction in survival rate was statistically significant.

Discussion

The micro-habitat use data confirm the hypothesis of an ontogenetic shift in Pterapogon kauderni micro­habitat put forward by Vagelli (2004). The data show that while Diadema micro­habitat is used by all life stages, recruits and small juveniles under 30 mm SL are most often as­sociated with sea anemones and hard corals are favoured by sub­adult (over 30 mm SL) and adult P. kauderni. Quan titative data from this study and qualitative data from long term observation of the introduced population in Palu Bay (unpublished data) indicate that recruit sur­vival is higher in sea anemones (with or without resident Amphiprion sp.) than in Diadema sea urchins.Like the clownfish (Amphiprion sp.), with which they

Table 1　P. kauderni gonad maturity parameter data

Fig. 4　Gut content composition of adult and sub­adult P. kauderni specimens examined

Ndobe et al.: The Banggai cardinalfish: an overview of local research248

often share an anemone, P. kauderni recruits are extremely vulnerable to predation by both adult P. kauderni and other carnivorous fish. Vagelli (2005) postulated that like-ly predators of P. kauderni included Crocodile-fish Cym­bacephalus beauforti (Platycephalidae), several species of  Lion-fish Pterois (Scorpaenidae), the grouper Epine­phelus merra (Serranidae), the Stonefish Synanceia hor­rida (Scorpaenidae), several species of moray eels of the

genus Gymnothorax, and Echidna nebulosa (Muraenidae) and the sea­snake Laticauda colubrina (Elapidae). Recent observations by the authors at Bone Baru, Banggai Island, include predation of recruits sheltering in Diadema urchins by adult P. kauderni,  stonefish  (Scorpaenidae), wrasses (Labridae), lizard fish (Cirrhitidae) and members of the Gobiidae Family. The observation near sea anem­ones of brooding males with larvae seemingly close to

Fig. 5　Gonad development by moon phase of female P. kauderni from Palu Bay: 5A Gonadosomatic Index (GSI); 5B Gonad maturity Index (GMI)

Fig. 6　P. kauderni gonads: 6A Female (with egg envelope); 6B Male

Fig. 7　Growth (7A) and Survival Rate (7B) of juvenile P. kauderni reared for 60 days at various salinities

Ndobe et al.: The Banggai cardinalfish: an overview of local research 249

their release as recruits indicates that male P. kauderni may choose to release their offspring near such protective micro­habitat. The data and observations suggest that micro­habitat availability is an important factor in P. kauderni population dynamics, with sea anemones being especially important for the survival of recruits, though further research is required to test this hypothesis.

At the time of the study, the authors warned that over exploitation of P. kauderni micro­habitat could have neg­ative effects on the population. Since the study was undertaken, significant reductions in P. kauderni popula­tions  have been  reported by fishermen  from Bone Baru (the main P. kauderni fishing village on Banggai Island) at sites where there has been a rapid expansion in seaweed farming, including at sites which are not P. kauderni fishing grounds. Seaweed farmers consuming benthic in-vertebrates in the areas around their farms, including large and increasing numbers of both sea anemones and sea urchins,  to replace fish which they often no longer have time to catch. The fact that both main micro­habitats (Diadema urchins and sea anemones) are exploited to the point of seriously reducing availability at an increasing number of sites is a worrying development which could become the main threat to the conservation of this endemic species. This is in contrast to the ornamental fishery and trade, previously considered the main threat to the species (e.g. Bruins et al. 2004, CITES 2007, Allen and Donaldson 2007, Vagelli 2008), in which there have been significant advances towards sustainability (Moore and Ndobe 2009 & 2011).

A previous study on P. kauderni diet in the wild (Vagelli and Erdmann 2002) found a similar preponderance of Crustacea to that found in this study. However Vagelli and Erdmann (2002) did not report Diadema or phytoplankton as components in P. kauderni diet. Phytoplankton do not form a large part of the observed diet and it has been suggested that they may be entrained with prey. Although Diadema are scarce at the Mamboro site, juvenile Dia­dema form part of the diet of P. kauderni, indicating a possible positive preference. Coppard and Campbell (2005) state that Diadematids provide a safe environment for many associated organisms, including cardinalfish. How-ever the benefits or effects on the sea urchin host are less well­known. It is hypothesized that the relationship be­

tween P. kauderni and Diadema sea urchins may be more complex than the simple use of the shelter provided by the Diadema urchins.

The difference in diet between adult and sub­adult P. kauderni, with adult diet consisting of a greater variety of organisms and less heavily dominated by copepods and other crustaceans than that of sub­adults, is unlikely to be related to increasing gape, as the additional items were not large. Vagelli and Erdmann (2002) stated that in their study P. kauderni food item size was not correlated with body size but did not mention any relationship between body size or life cycle phase and diet. The observed onto­genetic difference in diet could be related to reproductive maturity, specifically the nutrient requirements associated with the production of eggs and sperm.

Comparing gonad development observed in this study with the size of mouth­brooding males and observations on P. kauderni behaviour in other studies by the authors, both in  the laboratory and in  the wild,  indicate  that first maturity in P. kauderni may precede active reproduction. The data show that gonad maturation can occur at 30-35 mm SL, whereas mating behaviour and mouth­brood­ing have not been observed in wild P. kauderni populations at sizes below 40-42 mm SL. This could be due to com­petition for mates, which would be compatible with ob­servations by Kolm (2002) who found that female P. kauderni tend to prefer (and can even produce larger num­bers of eggs for) larger males.

A relationship between the lunar cycle and P. kauderni breeding was suggested by Vagelli and Volpedo (2004), with a major peak around the full moon and a minor peak around the new moon, based on P. kauderni specimens from the Banggai Archipelago. However specimens from one site (Tempaus, a site with environmental conditions somewhat different for most of the endemic distribution) did  not  fit  this  pattern.  In  Palu Bay  only  one  peak was observed, around the full moon period. This has since been  confirmed  by  many  subsequent  observations  of breed ing behaviour in the Palu Bay introduced population and recent observations at Bone Baru, Banggai Island indicate a similar pattern at this site. Pair formation and the defence of breeding territories have been observed shortly before the full moon (second quarter, waxing moon), with many breeding males and recruits generally

Ndobe et al.: The Banggai cardinalfish: an overview of local research250

visible in the third quarter (waning moon). It is considered likely that P. kauderni reproduction is linked to the lunar cycle but that the timing of spawning peaks is influenced by site-specific factors which could be elucidated through studies at multiple sites with different environmental characteristics.According  to Bone and Moore  (2008) most fish have 

bi­lobed (paired) gonads and mechanisms for determining the  sex  of  individual  fish,  which  vary  greatly  between species, and can include genetic, environmental, and social factors. Fishelson and Gon (2008) studied the egg envelopes of 30 species of mouth­brooding Apogonidae and state that all have bi­lobed gonads. The nearest known relative to P. kauderni (Pterapogon or Quinca mirifica) is known to have standard bi­lobed gonads (Vagelli, 2009). The authors have not found any references regarding P. kauderni gonad morphology, mentioning mechanisms by which the sex of individual P. kauderni might be deter­mined or indeed indicating the existence of other non bi­lobed apogonid species. Further research into P. kauderni gonad morphology and development is currently being planned.

The sample observed in this study was too small to conclude that the unequal sex ratio observed is a general characteristic, even though there were more males than females in the randomly selected sample in every moon phase. Vagelli and Volpedo (2004) concluded that P. kauderni had a sex ratio of one. However although the difference  was  not  significant  using  standard  statistical tests, in that study also there was an excess of males over females in the sample they obtained. Given that repro duc­tive output is limited by male capacity to mouth­brood, a higher number of males would logically result in greater population resilience and capacity for growth. Thus sex ratio is an important parameter for the management of P. kauderni populations. If sex is environmentally deter­mined, the sex ratio could vary in both time and space. Further research is considered necessary on the sex ratio and mechanisms of sex determination in P. kauderni.

P. kauderni juveniles exhibited a relatively wide salinity tolerance (euryhaline range), adapting to salinity from 24-35 ppt. The juvenile P. kauderni were able to withstand much lower salinity levels for short periods of time and even at the lowest salinity (16 ppt) treatment, with a sur-

vival rate as low as 27% over 60 days, there was no sig-nificant difference in survival rate or growth for the first 10 days. This could be related to the inherent variability in water quality in shallow (less than 5 m deep) of the coastal waters which comprise the endemic habitat, for example, salinity fluctuations related to rainfall and run-off as well as evaporation. This tolerance has implications for the culture and transport of P. kauderni as well as for popu­lation resilience and the ability to establish viable popu­lations when introduced to sites outside the endemic range. It is possible that even lower salinity could be tolerated for short periods of time. A local ornamental fish trader claims to have adapted P. kauderni to fresh water conditions, however the authors have not been able to substantiate this and an attempt to replicate the process by students under controlled conditions resulted in 100% mortality.

The research undertaken by authors was designed to support the ongoing BCF Action Plan. In this context, the authors made a number of recommendations. Firstly, that habitat conservation should be a major priority for P. kauderni management, especially the three key micro­habitats (Diadema urchins, sea anemones and hard corals), including  finding  alternatives  for  human  consumption. Secondly, that fishing activities should be timed to avoid peaks in breeding activity, at least at sites where the timing is  known.  Thirdly,  the  findings  highlight  the  need  for further research to gain a more comprehensive knowledge of P. kauderni biology and ecology, including many aspects of reproductive biology and population dynamics. In particular there is a need to establish the factors de­termining population resilience to natural and anthro­pogenic pressures and develop a P. kauderni population dynamics model, and combine this with socio­economic data in order to plan and implement ecologically and socially sustainable management.

Acknowledgments

The authors wish to thank all who have in any way contributed to the research presented and the production of this paper. In particular, funding for some research was provided by two grants from DIKTI, the Indonesian

Ndobe et al.: The Banggai cardinalfish: an overview of local research 251

Higher Education Authority and attendance at the 2nd APCRS was supported by a grant from the BP Conservation Leaders Programme and assistance from the APCRS com mittee. Special acknowledgment is due to all who took part in or otherwise supported any aspect of the research presented, especially students and colleagues from Tadulako University (UNTAD), the Sekolah Tinggi Perikanan dan Kelautan (STPL), LP3L Talinti and the Banggai Cardinalfish Centre (BCFC).

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