Japanese Journal of Ichthyology 魚 類 学 雑 誌 Vol. 33, No. 3 1986 33巻3号1986年

Courtship and Spawning of the Hawkfish of sea anenomes (Parasicyonis spp.) were clustered falco at Miyake-jima, on the seaward margin of the boulder, directly Japan beneath the outcropping. The adjacent sub stratum consisted of five smaller boulders (ca. 0.5-2.0 m high), mixed volcanic rubble, sand, and Terry J. Donaldson various algae. A slight to moderate current, (Received December 28, 1985) made stronger occasionally by the effects of passing swells generated by storms well offshore, The hawkfish Cirrhitichthys falco Randall flowed seaward past the site. Water temperatures (Cirrhitidae) occurs from Japan (Masuda et al., ranged from 23 to 26℃. 1975) south to the Philippines (Randall , 1963), the Observations were made using SCUBA, and Marianas (Myers and Shepard, 1980), Pohnpei data were recorded on plastic slates and by (Gawel, unpubl. ms.), Fiji (Springer, 1982) to underwater photography. Two dives were made northeastern Australia (Russell, 1983) as far as daily, usually between 10: 30-14: 00h and 17: 45- Sydney (R. E. Thresher, pers. comm.). Although 19 : 00h. The first dive consisted of a population this species occurs at Miyake-jima, lzu Islands, census and observations of habitat utilization, Japan(34°05′N,139°30′E) it is rare and may territorial boundaries, and aggressive behavior, often be mistaken for C. aprinus (Bleeker). while the second dive was devoted to observations Courtship and spawning behavior of cirrhitid of courtship and spawning. has been largely limited to aquarium ob servations. Lobel (1974), Takeshita (1975) and Tanaka et al. (1985a) reported on courtship and Results spawning of (Bleeker); Suzuki et al. (1980) reported spawning behavior of Cir Social organization and aggressive behavior. Two , a male (designated as Ml, ca. 80 mm rhitichthys aureus (Temminck et Schlegel); Ta SL) and a female (F1, ca. 70 mm SL) were ob naka et al. (1985b) reported spawning behavior of served near and at the rendezvous site. No hubbardi (Schultz). other conspecifics were present in the area. The Field studies of cirrhitid courtship and spawning female occupied a small territory (ca. 3 m2) im behavior have begun to appear . Thresher (1984) mediately beneath the outcropping on the shore briefly mentioned courtship of ward side of the boulder. She was quite secretive arcatus (Cuvier) from One Tree Island , Great and rarely left the shelter of a small hole located Barrier Reef and has also provided a detailed in this area. The male foraged over a territory, description of courtship and spawning of Cir ca. 25 m2, adjacent to and including the ren rhitichthys oxycephalus (Bleeker) occurring in the dezvous site. No active territorial defense against Gulf of California. Here I report courtship and conspecifics was observed. The male did defend spawning behavior of a congener , Cirrhitichthys parts his territory against some interspecific in fako, at Miyake-jima. truders. Many territorial interactions were with a neighboring congener, a large male (ca. 90 mm Study area and methods SL) C. aprinus, which occurred at the rendezvous site and at a common territorial boundary shared Courtship and spawning behavior of C. falco with this species. Defense was particularly pro were studied on seven consecutive days between nounced just prior to and after sunset when court 6-12 August 1985 in Igaya Bay. All observa ship between M1 and F1 was about to commence. tions were made at a single site located ca. 100 m Males of both species interacted aggressively with NE of the pier at Igaya Port at a depth of 10 m. one another but not with the female C. falco. The courtship and spawning site, henceforth re Courtship and spawning. Five distinct motor ferred to as the rendezvous site (Moyer , 1980, patterns were recognized during male-female in 1984), consisted of a rocky outcropping located teractions: 1) Parallel Rest, 2) Nudging, 3) Circl atop a large boulder (ca. 2.5 m high) immediately ing, 4) Hopping and 5) Spawning. Circling and adjacent to an abandoned pipeline . A number Parallel Rest were also seen in interactions be-

―329― 魚類学雑誌 Japan. J. Ichthyol. 33(3), 1986 tween male C. falco and C. aprinus. Motor their snouts downward (Fig. 1E) from the parallel patterns are described as follows : rest position, and released a cloud of gametes 1) Parallel Rest. The male and female aligned before quickly returning to the substratum. themselves on the substratum in a position parallel Courtship and spawning events involved only to one another (Fig. 1A). Occasionally, the male M1 and F1 although the male C. aprinus attempted was positioned slightly behind the female. This courtship with F 1 on a number of occasions but motor pattern often preceeded the performance was always driven off by M1 . Time of courtship of other motor patterns and always preceeded commencement ranged from one minute before spawning. sunset to 20 minutes past sunset over a period of 2) Nudging. During Nudging the male placed seven days and lasted between 2-16 minutes before his snout against the flank of the female at a point spawning occurred (Table 1). Typically, either directly beneath the soft dorsal fin (Fig. 1B) M1 or F 1 arrived at the rendezvous site first and while both the male and female remained mo was joined immediately by the other. On one tionless. The behavior was seen soon after pair occasion M1 was observed to herd F 1 towards formation and was often repeated during a the rendezvous site from a position directly below courtship encounter. This motor pattern is the site. similar to that reported for C. oxycephalus by Once at the rendezvous site the pair first ex Thresher (1984). hibited the Parallel Rest motor pattern for 5-20 s 3) Circling. The male positioned his snout and then the male nudged the female. Hopping near the caudal fin of the female and vice versa; and Circling followed with intervals of Parallel together they swam in a circular fashion (Fig. Rest and Nudging and ended in the Parallel Rest 1C) around the rendezvous site and made frequent for a period of 10-40 s. The spawning ascent stops to Parallel Rest. This motor pattern fre immediately followed the last Parallel Rest display. quently occurred while the male and female were Spawning times varied between 4-22 minutes past engaged in Hopping. sunset (Table 1). After the release of gametes, 4) Hopping. The male and female engaged the small white cloud was visible for 3-9 s. Egg in short swimming bouts, resembling hopping, predation was not observed. around the rendezvous site. The male was posi After returning to the substratum the male tioned slightly behind the female as they swam attempted to court the female again on five of (Fig. 1D). Hopping was often preceeded and seven evenings when spawning was observed. followed by Parallel Rest displays. This motor The male assumed the Parallel Rest position and pattern was also reported for C. oxycephalus by nudged the female. However, this second attempt Thresher (1984). at courtship was never successful; either the female 5) Spawning. After the last Parallel Rest returned to her shelter beneath the rendezvous display the male and female ascended into the site or the courtship behavior was interrupted by water column to a height of 20-50 cm, pointed the approach of the male C. aprinus or other

Table 1. Temporal patterns of courtship and spawning of the hawkfish Cirrhitichthys falco at Miyake-jima, Japan.

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A B C

D E

Fig. 1. Motor patterns observed during courtship and spawning of the hawkfish Cirrhitichthys falco at Miyake-jima, Japan. A) Parallel Rest, B) Nudging, C) Circling, D) Hopping and E) Spawning. (Male denoted by M; female denoted by F).

species of fishes resident in the area. rhitidae. The occurence of a male-female pair of Interspecific interactions. Courtship behavior C. falco at the spawning site was consistent with was often interrupted by the presence of the male observations of this species made at Guam, C. aprinus or other species of fishes, resulting in Mariana Islands although social groups of up to variable durations in time before spawning oc six individuals have also been seen (pers. obs.). curred (Table 1). The male C. aprinus interrupted Thresher (1984) reported social groups of one male courtship both before and after the spawning; and up to seven females for C. oxycephalus in the interruption took the form of swift charges at M 1 Gulf of California. I have seen male-female pairs and attempted courtship with F1 before M1 was only of the following other species of cirrhitids : able to drive the male C. aprinus away. Court O.typus at Pohnpei, Caroline Islands; Paracir ship motor patterns displayed by the male C. rhites hemistictus (Gunther) at Pohnpei and aprinus included the Parallel Rest and Hopping. Guam; P. forsteri (Schneider) at Pohnpei and Other species of fishes in the area interrupted Guam; Castelnau at Guam. courtship both before and after M1 and F1 However, N. armatus, P. arcatus, P. forsteri and spawned (Table 1). Courting male and female P. hemistictus have been observed also in groups Anthias (Franzia) squamipinnis (Peters), foraging of one male and up to four females at Guam. Epinephelus fasciatus (Forsskal) and territorial J. T. Moyer (pers. comm.) reported sightings of Stegastes altus (Okada et Ikeda) all interrupted two C. aprinus social groups at Igaya Bay, Miyake- C. falco during the first bout of courtship prior jima during 1984-85. Each social group con to spawning; an aggregation of Pempheris sisted of one male and two females. Both social oualensis Cuvier interrupted post-courtship dis units disappeared during a typhoon in June, 1985. plays after spawning on one occasion. This same typhoon may account for the attempts made by the male C. aprinus at courting the Discussion female C. falco; the former may have lost his mate(s) by the typhoon. Moyer (1981) observed Social organization and mating system in Cir interspecific spawning between a male Centropyge

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shepardi Randall et Yasuda and a female C. floating eggs to escape predation from bottom bispinosus (Gunther) at Guam, where the latter dwelling organisms by their release into the water species is rare and the former is common. Moyer column above the substratum. The height of hypothesized that such interspecific spawnings the release may be constrained by the risk of usually involve females of rare species and males predation on the spawning adults (Thresher, of common species. Such spawnings may occur 1984). Emery (1973), Robertson and Hoffman as a result of constraints imposed by energy ex (1977), Moyer and Yogo (1982) and Thresher penditure in the production of eggs versus sperm; (1984) reported that predators often attacked females should engage in interspecific spawnings spawning individuals; the probability of such only when male conspecifics are absent whereas attacks may be minimized by rapid ascents to males can afford to be less discriminating in minimal heights that would allow spawning adults choosing mates. to avoid the risk of predation while conveying an Thresher (1984) considered C. oxycephalus to anti-predation advantage to the free-floating eggs possess a mating system of male-dominated social released during the ascent (Thresher, 1984). A ly controlled hermaphroditism. Kobayashi et al . test of this hypothesis on small site-attached (1985) recently reported the existence of pro species seems warranted. togynous hermaphroditism in C. falco and in C. aureus, C. aprinus and Cirrhitops hubbardi as well. Acknowledgments Donaldson (unpubl. ms.) has recently described the mating and social systems of P. arcatus, P. I wish to thank J. T. Moyer, J. E. Randall, forsteri, P. hemistictus, and N. armatus. P. L. Colin, L. J. Bell-Colin, D. Y. Shapiro, J. M. Male defense of both the rendezvous site and Fitzsimons, M. C. Wilkins and S. de C. Wilkins the female was indicated in C. falco, but the rela for their advice, comments and assistance. G. B. tive importance of defending one over the other Constantino kindly prepared Fig. 1. I also thank is uncertain. J. T. Moyer (pers. comm .) reported the Tatsuo Tanaka Memorial Biological Station that males of C. aprinus at Miyake-jima travelled for its hospitality. This work was made possible over "well-known" paths in given areas but did through the generous support of the Japan Airlines not engage in intraspecific territorial defense of 50th 747 Asian Studies Scholarship and the LSU these areas although they did defend females from Museum of Natural Science. This is contribution male conspecifics. no. 64 of the Tatsuo Tanaka Memorial Biological Interspecific comparisons of courtship and spawn Station. ing. Four of the C. falco courtship and spawning motor patterns observed at Miyake-jima are rela Literature cited tively consistent with those observed for C. oxycephalus in the Gulf of California (Thresher, Emery, A. R. 1973. Comparative ecology and func tional osteology of fourteen species of damselfish 1984). These patterns are Parallel Resting , Nudging, Hopping and the spawning ascent. (Pisces: Pomacentridae) at Alligator Reef, Florida Keys. Bull. Mar. Sci., 23: 649-770. Male C. oxycephalus approached females at a Kobayashi, K., K. Suzuki, T. Suzuki and S. Mishina. single spawning site and, in succession, nudged, 1985. Hermaphroditism in the four Japanese hopped, and rested before executing the spawning cirrhitid fishes. Advance Abstracts for the 18th ascent. Two of the motor patterns, Nudging and Annual Meeting of the Ichthyological Society of the spawning ascent, seem common of pelagic Japan. No. 18. (In Japanese.) spawning reef fishes in general. Males of C. Lobel, P. S. 1974. Sea spawnings-hawkfish. Octo aprinus apparently nudge females by lying over pus, 1(7): 23. their bodies (J. T. Moyer, pers. comm.); I have Masuda, H., C. Araga and T. Yoshino. 1975. Coast observed males of N. armatus performing a similar al fishes of southern Japan. Tokai University Press, Tokyo, 382 pp., 151 pls. pattern during courtship. The significance of the Moyer, J. T. 1980. The mating strategy of the Nudging motor pattern is uncertain, although thornback cowfish, Lactoria fornasini. Anima, (91): Thresher (1984) speculated that it served to 50-55. (In Japanese.) stimulate egg movement in the female prior to Moyer, J. T. 1981. Interspecific spawning of the spawning. The spawning ascent allows free pygmy angelfishes Centropyge shepardi and C.

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bispinosus at Guam. Micronesica, 17: 119-124. Takeshita, G. Y. 1975. Long-snouted hawkfish. Moyer, J. T. 1984. Social organization and re Mar. Aquar., 6(6): 27-31. productive behavior of ostraciid fishes from Japan Tanaka, Y., Y. Shiobara, T. Ohyama, I. Ozaki and and the western Atlantic Ocean. J. Ethol., 2: 85- M. Moriya. 1985a. Spawning behaviour, eggs 98. and larvae of the hawkfish, Oxycirrhites typus, in Moyer, J. T. and Y. Yogo. 1982. The lek-like an aquarium. Advance Abstracts for the 18th mating system of Halichoeres melanochir (Pisces : Annual Meeting of the Ichthyological Society of Labridae) at Miyake-jima, Japan. Z. Tierpsychol., Japan, No. 57. (In Japanese.) 60: 209-226. Tanaka, Y., Y. Shiobara, M. Hayashi, T. Furukawa Myers, R. F. and J. W. Shepard. 1980. New records and M. Hattori. 1985b. Spawning behaviour, of fishes from Guam with notes on the ichthyofauna eggs and larvae of the hawkfish, Cirrhitops hubbardi, of the southern Marianas. Micronesica, 16: 305- in an aquarium. Advance Abstracts of the 18th 347. Annual Meeting of the Ichthyological Society of Randall, J. E. 1963. Review of the hawkfishes Japan, No. 58. (Family Cirrhitidae). Proc. U.S. Natn. Mus., 114 Thresher, R. E. 1984. Reproduction in reef fishes. (3472): 389-451, pls. 1-16. TFH. Publ., Neptune City, 339 pp. Robertson, D. R. and S. Hoffman. 1977. The roles of female mate choice and predation in the mating (Section of Ichthyology, Museum of Natural Science systems of some tropical labroid fishes. Z. Tier and Department of Zoology and Physiology, Louisiana psychol., 45; 289-320. State University, Baton Rouge, Louisiana 70893, Russell, B. C. 1983. Annotated checklist of the USA). reef fishes in the Capricorn-Bunker Group, Great Barrier Reef, Australia. Great Barrier Reef 三 宅 島 に お け る サ ラ サ ゴ ン ベ の 求 愛 ・産 卵 行 動 Marine Park Authority, Townsville, Queensland, 184 pp., 49 pls. Terry J. Donaldson Springer, V. G. 1982. Pacific Plate biogeography, 1985年8月,伊 豆 諸 島 の 三 宅 島 に お い て サ ラ サ ゴ ン ベ(ゴ ン ベ 科)の 産 卵 行 動 が は じ め て 観 察 さ れ た.1週 with special reference to shorefishes. Smithson . Contr. Zool., 367: i-iv +1-182. 間 の 調 査 期 間 中,雄(体 長8cm)と 雌(7cm)の ペ ア Suzuki, K., Y. Tanaka, Y. Suzuki and H. Tanaka. が 産 卵 場 所 と な る 岩 を 中 心 と し て 定 住 し て い た.毎 日, 1980. Spawning behavior and early life history of 日 没 直 後 に な る と,ペ ア は 定 っ た 岩 の 上 で 一 連 の 求 愛 行 the hawkfish, Cirrhitichthys aureus, in the aquarium . 動 を 始 め,3-16分 後 に20-50cm急 上 昇 し て 放 卵 放 精 Advance Abstracts for the 13th Annual Meeting of し た.産 卵 前 後 に は,同 属 の ミ ナ ミ ゴ ン ベ の 雄 が こ の 雌 the Ichthyological Society of Japan, No. 40. (In に 求 愛 す る こ と や,そ の 他 の 魚 種 に よ る 干 渉 も し ば し ば Japanese.) 観 察 さ れ た.本 種 の 社 会 構 造 ・婚 姻 組 織 お よ び 求 愛 ・産 卵 行 動 を 他 の ゴ ン ベ 科 の 魚 類 と 比 較 し て 考 察 し た.

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