Batesian Mimicry Between a Cardinalfish (Apogonidae) and a Venomous Scorpionfish (Scorpaenidae) from the Philippine Islands!

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Batesian Mimicry between a Cardinalfish (Apogonidae) and a Venomous Scorpionfish (Scorpaenidae) from the Philippine Islands!

JEFFR EY A. SEIGEL 2 and THOMAS A. ADAMSON3

ABSTRACT: An apparent case of Batesian mimicry involving the cardinalfish Fowleria sp. and the venomous scorpionfish Scorpaenodes guam ensis is de scribed . Classical definitions of Batesian systems require that the mimic be less common than the model. The present data do not suggest a classical Batesian mimetic system. However, if the model is con sidered to be the generalized scorpaenid morph characteristic of S. guamensis and other Philippine scor paenids, then the mimetic association between Fowleria sp. and S corpaenodes guamensis is a classical Batesian system.

OF THE REPORTED OBSERVATIONS of inter Ru ssel, Allen, and Lubbock (1976) noted specific mimicry in fishes (e.g., Whitley 1935, that most reported cases ofinterspecific mim Randall and Randall 1960, Tyler 1966,Wickler icry in fishes can be classified as either 1968, Hobson 1969, Starck 1969, Losey 1972, Batesian, Mullerian, or aggressive mimicry. Springer and Smith-Vaniz 1972, Russel, Batesian mimicry typically includes three ele Allen, and Lubbock 1976, Thresher 1978, ments: one or more predator species; a model Ormond 1980), there is only Whitle y's (1935) that is dangerous or unpalatable to the pre record of a mimetic association involving a dator; and a mimic species that obtains some species of venomous scorpaenid and a non protection from predation by resembling the venomous mimic. Whitely described several model (Nur 1970, Greene and McDiarmid specimens ofthe serranid Centrogenys vaigen 1981). sis and the scorpaenid Sebastapistes bynoensis We report a new case of apparent Batesian laotale collected from North Queensland, mimicry between a species of apogonid, Australia, noting that the former, a percoid, Fowleria Sp.,4 and a venomous scorpaenid, was a mimic of the latter in both form and Scorpa enodes guamensis, from the Philippine coloration. He remarked that scorpaenoids Islands. Material is deposited in the Natural are "sluggish creatures with venomous spines History Museum of Los Angeles County on the head or in the dorsal fin, so that the (LACM). All measurements are standard length harmless Centrogenys may derive some for (SL). tuitous benefit from resembling them ." In Ju ly of 1981, while diving at Puerto Galera, Oriental Mindoro, Republic of the 1 Field work in the Philippine Islands was supported by Philippines, one of us (Seigel) observed a Natu ral History Museum of Los Angeles County Foundation Travel Grant No. 104. Manu script accepted mimetic association involving the cardinalfish 5 November 1982. Fowleria sp. and the scorpionfish Scorpaenodes 2 Natural History Museum of Los Angeles County, guamensis. Two stations (PI81-36, 37, Table Section of Fishes, 900 Exposition Boulevard , Los 1) were particularly note worthy, for in 5 or 6 Angeles, California 90007. separate localities, it was observed that when 3 Natural History Museum of Los Angeles County, Section of Fishes, 900 Exposition Boulevard, Los ichthyocide was applied to small caves or to Angeles, California 90007. Present address (TA) : interstices in large rock boulders, a small Adamson Philippines Inc., 51 E. Colorado Boulevard, S. guamensis (sizes ranged 15.0-68.0mm SL) Pasadena, California 91105. swam away from the rocks (usually 5 to 10 4 Identification is tentativ e, since the genus Fowleria is in need of revision. There are seven nominal species min after application of ichthyocide) closely (Fraser 1972) of which F. variegat us most closely re followed by at least one Fowleria of nearl y sembles the Fowleria figured in this paper. the same size; in some instances mor e than 75 76 PACIFIC SCIENCE, Volume 37, January 1983

FIGURE I. Scorpaenodes guamensis (A, D, E, H ) and its mimic Fowleria sp. (B, C, F, G). Lateral view of A, LACM 42490-56, I of 3, 39.0 mm; B, LACM 42490-51, I of lO, 34.0mm; C, LACM 42490-51, I of 10, 27.0mm; D, LACM 42490-56, 1 of 3, 23.0 rnm; E, LACM 42491-52, 1 of 6, 33.0mm; F, LACM 42491-49, 1 of 5, 33.0 mm; G, LACM 42491-49, 1 of 5, 26.0mm; H , LACM 42491-52, I of 5, 15.0mm.

TAB L E 1 caudal fins are variably banded, thepelvic fins PHILIPPINE ISLAND STATIONSTHAT YIELDED Fowleria dark, and there are two or more oblique bars SP. AND S. guamensis INCLUDING NUMB ER OF EACH extending ventrally and posteriorly from the SPECIES rim ofthe orbit to the maxillary and the pos terior margin of the preopercle, respectively STATION LOCALITY Fowleria SP. S. guamensis (Figure I). Body shape is remarkably similar in the two species . Both appear as rather PI81-2 Hundred Island s 10 2 PI81-7 Hun dred Islands 6 3 robust fishes, and the dorsal and ventral pro PI81-13 Lingayen Gulf 6 I files ofthe head and body are nearly identical PI81-35 Puerto Galera 5 I (Figure I). The mouth is upturned at an angle PI81-36 Puerto Galera 10 3 of approximately 25-30° and the eye is large PI81-37 Puerto Gale ra 5 6 (30-40 percent head length). The Scorpaenoidei is reno wned for including the world's mo st venomous fishes (Halstead one Fowleria was taken along with a small and Mitchell 1963, Halstead 1970, Nelson S. guam ensis. 1976). Halstead and Mitchell (1963) recog Both species are strikingly similar in colora nized the genus Scorpaenodes as a venomous tion, the live-color patterns con sisting ofdark scorpionfish, a statement sub sequently corro reddish brown mottled bands alterna ting with borated by Halstead (1970), who listed two light-colored bands flecked with small reddish venomous species of Scorpa enodes, S. gua brown spots (Figure I). The dorsal, anal, and mensis and S. parvipinnis. There are at least

;ng*ib:tid= Mimicry in Philippine Fishes-SEIGEL AND ADAMSON 77

FIGURE 2. Morphologically similar Philippine scorpaenids. Lateral view of A , Scorpa enodes guam ensis, LACM 42491-52, 1 of 6,45.0 mm; E, Scorpaenodes scabra, LACM 42456-41,67.0 mm ; C, S corpaenopsis cirrhosa , LACM 42479 37,61 .0 mm ; D, Scorpaenopsis diabolus, LACM 35977-30, 36.0 mm ; E, Sebastapistes tristis, LACM 42467-42, 34.0 mm; F, Parascorpaena aurita , LACM 42461-58, 72.0 mm. seven common Philippine scorpaenids that overlapping. S. guamensts IS found along resemble Scorpa enodes guamensis in overall rocky and coral reefs throughout the tropical body shape and coloration: Scorpaenodes Indo-west Pacific (Eschmeyer 1969, Allen scabra, Hypomacrus albaiensis, Scorpaenopsis and Steene 1979) and Fowleria (with seven cirrhosa, Scorpaenopsis diabolus, Sebastapistes nominal species, see footnote 4) is widely dis tristis, and Parascorpaena aurita (Figure 2, tributed in the Indo-west Pacific from east Hypomacrus albaiensis not figured). Most of Africa to Samoa, the Philippine Islands, and these are known to be venomous (Halstead northward to Japan (Fraser 1972, Masuda, 1970). Araga, and Yoshino 1975,Yoshino and Nishi There are no known venomous cardinal jima 1981). fishes. The Apogonidae contains about 200 The present observations are based on ma species representing 24 to 26 diverse genera, terial collected during June and July 1981 most of which are small (less than 100mm), from 39 stations in 3 separate regions of the secretive coral reef fishes (Fraser 1972). Philippine Islands: Hundred Islands and adja Randall and Randall (1960) noted that cent areas, Lingayen Gulf, northwest Luzon ; evidence for mimicry should follow other Honda Bayand adjacent areas, and Sombrero lines in addition to the similarity of the two Island, Palawan; and Puerto Galera, Oriental species in form and color: "a mimic occupies Mindoro. A review of species lists from each the same habitat as the model, and its distri station revealed that Fowleria sp. and bution frequently coincides with that of the S. guamensis were collected in 6 of 39 stations model." The geographic ranges of Fowleria (3stations in Hundred IslandsjLingayen Gulf; sp. and Scorpaenodes guamensis are broadly 3 stations in Puerto Galera), and except for 78 PACIFIC SCIENCE, Volume 37, January 1983

I station in Honda Bay, Palawan , neither spe specialized mimics, tho se that are most strik cies was taken in a collection without the ing to the observer. The description of a new other. caseof Batesian mimicry involving a venomous scorpionfish and a nonvenomous cardinalfish DISCUSSION presents evidence that mimi cry may in fact be a relatively widespread, though often un The numbers of both species collected observed or unrecognized, phenomenon in at each station do not suggest the classical marine fishes. Batesian complex in which the mimic is less common than the model (Table I). The pre sent data indicate that in collections made at ACKNOWLEDGM ENTS stations where Fowleria sp. and S. guamensis were present , the mimic-to-model rat ios We thank W. N . Eschmeyer for identifica ranged from I : I to 6 : I (Table I). Previous tion of scorpaenids, R. J. Lavenberg, C. C. authors have discussed mimic-to-model Swift, and J. W. Wright for review of the ratios in Batesian systems, noting that there manuscript, and B. N. Wh ite for helpful are numerous situa tions in which Bate sian comments. We also thank M. deGuzman for mimics occur in equal or greater numbers her help while collecting .in Lingayen Gulf. relative to models (Fisher 1930, Springer and Photographs were taken by J. DeLeon and Smith-Vaniz 1972, Ru ssel, Allen , and Lub R. Meier (LACM). bock 1976, Greene and M cDiarmid 1981). Innate avoidance, as described by Springer and Smith-Vaniz (and subsequently by Smith LITERATURE CITED 1975 and Greene and McDiarmid 1981) is an individually variable, genetically based re ALL EN, G. R., and R. C. STEENE. 1979. The sponse whereby predators innately recognize fishes of Christmas Island, Indian Ocean. a particular morphology or behavior as be Au st. Nat. Parks and Wildlife ServoSpec. longing to an unpalatable prey species. By Pub. No . 2. mimicking Sco rpaenodes guamensis, the non ESCHMEYER, W. N . 1969. A new scorpionfish venomous cardinalfish Fowleria sp. may gain ofthe genus Scorpaenodes and S. mu ciparus protection from predation becau se it closely (Alcock) from the Indi an Ocean, with com resembles a generalized, venomous scorpaenid ments on the limits of the genus. Occ. Pap. morph innately avoided by potential preda Calif. Acad. Sci. No. 76. tors. There are a number of very similar ven FISH ER, R. A. 1930. The genetical theory of omous scorpaenids (see above and Figure 2) natural selection. Clarendon Press, Oxford. that pro vide, along with S. guamensis, a F RASER, T. H . 1972. Comparative osteology "model morph" for the cardinalfish Fowleria of the shallow water cardinal fishes (Perci sp. It is possible that the mimic, Fowleria sp., formes: Apogonidae) with reference to the is in fact less abundant than its model, if the systematics and evolution of the fam ily. model is considered to be the generalized, ven leh. Bull. J. L. B. Smith Inst. Ich. 34 : 1 omous scorpaenid morphology characteristic 105. of S. guamensis and many other Philippine GREENE, H. W., and R. W. M cDIARMID. scor pionfishes. 1981. Coral snake mimicry: doe s it exist? Greene and McDiarmid (1981) stated that Science 213 : 1207-1212. Batesian and Mullerian mimicry are rare HALSTEAD, B. W. 1970. Poisonous and ven phenomena among vertebrates. Conversely, omous marine animals ofthe world. vol. 3, Ru ssel, Allen , and Lubbock (1976) noted that Vertebrates. U .S. Govt. Printing Office, mimicry, at least among marine fishes, may Washington, D.C. actually be relativel y general and widespread ; HALSTEAD, B. W., and L. R. MITCHELL. 1963. they stated that previou s description s ofmim A review of the venomous fishes of the Pa icry centered around the more spectacularand cific area. Pages 173- 202 in H. L. Keegan

i ' t Mimicry in Philippine Fishes-SEIGEL AND ADAMSON 79

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