Istiophoridae, Xiphiidae, and Xiphiorhynchidae)

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Reprinted from: Stroud. Richard H. (ed.). 1990. Planning the Future of Billfishes. Part 2. COlltrib~ted Papers. Proceedings of the Second Intem.~tlonal Billfish Symposium, Kailua-Kana, HawaII, August 1-5, 1988.

A Paleontological Review of Three Billfish Families (Istiophoridae, Xiphiidae, and Xiphiorhynchidae)

Harry L. Fierstine

In 1974, I summarized the osteological and the taxonomic nomenclature of Nakamura paleontological knowledge of billfishes and (1983), with the exception that I agree with Robins suggested some areas of research to aid future and de Sylva (1960) that both the sailfish and paleontological studies (Fierstine 1974). My blue marlin exist as single world-wide species, suggestions included: (l) a comparative osteo­ and not as separate Atlantic Ocean and Pacific logical study, especially of the rostrum, in order Ocean forms. to obtain more accurate identifications of fos­ Family Xiphiidae silized material; (2) a functional study of the Xiphias Linnaeus feeding mechanism, in order to explain observed Xiphias gladius Linnaeus, 1758 ­ interfamilial differences in the skull; (3) a func­ Swordfish tional study of the locomotor apparatus in order Family Istiophoridae to elucidate inter- and intrafamilial differences Istiophorus Lacepede in vertebrae, body shape and fin morphology, Istiophorus platypterus (Shaw and and (4) a more thorough study of those extinct Nodder, 1792) - Sailfish families (Blochiidae and Paleorhynchidae) and Makaira Lacepede genera (Acestrus and Brachyrhynchus) having Makaira indica (Cuvier, 1832) - Black uncertain affinities to other billfishes. marlin Since 1974, some of the foregoing items, as Makaira nigricans Lacepede, 1802­ well as some not listed, have been studied with Blue marlin varying degrees of thoroughness. In the most Tetrapturus Rafinesque comprehensive treatise, Schultz (1987) investi­ Tetrapturus albidus Poey, 1860­ gated all extant and extinct billfishes and White marlin developed a new taxonomic arrangement and Tetrapturus angustirostris Tanaka, 1915 evolutionary scheme based primarily on structure - Shortbill spearfish of the rostrum. The present study reviews the Tetrapturus audax (Philippi, 1887)­ published accounts, presents some new informa­ Striped marlin tion, and summarizes the relevant osteological Tetrapturus belone Rafinesque, 1810­ and current paleontological knowledge of three Mediterranean spearfish billfish families (lstiophoridae, Xiphiidae, and Tetrapturus georgei Lowe, 1840­ Xiphiorhynchidae) . Roundscale spearfish Tetrapturus pfluegeri Robins and de Systematics of Recent Forms Sylva, 1963 - Longbill spearfish Al'.hough Schultz (1987) rearranged the extant billfishes into three families (lstiophoridae), Relevant Osteology of Recent Forms Tetrapturidae, and Xiphiidae) and six genera Skull (Istiophorus, Makaira, Marlina, Pseudohis­ General account and terminology. I will follow tiophorus, Tetrapturus, andXiphias), I will follow the lead of other billfish scientists (e.g., Nakamura

11 12 PLANNING THE FUTURE OF BILLFISHES

1983) and use the osteological terminology of three istiophorid genera can be distinguished by Gregory (1933) and Gregory and Conrad (1937) the shape of their rostra at the rostral mid-point where possible even though it may not conform (Fig. 2). The distribution of denticles (Table I) to modern comparative developmental studies and relative size of the paired nutrient canals (Jollie 1986). Myer et al (in press) and Schultz show interspecific patterns. (1987) independently discovered that the Schultz (1987) noticed that the denticles along istiophorid rostrum contained a new paired ele­ the ventral midline of the rostrum of the shortbill ment, the prenasal, which had mistakenly been spearfish and Mediterranean spearfish are not considered part of the premaxillary bone by housed in alveoli and detach when the rostra are other workers. prepared into skeletons. He separated all spear­ Rostrum. The rostrum (or bill) is defined as fishes (T. angustirostris, T. belone, T. georgei, that part of the snout which extends anteriorly and T. pfluegeri) from the other billfishes from the anterior border of the bony orbit to primarily on the basis of the edentulous zone. terminate in a projection beyond the lower jaw There was no mention of intraspecific or in­ (Fierstine 1978). The Istiophoridae and Xiphiidae terspecific variation of the character. have rostra with different morphologies. Poplin et al (1976) described the microscopic The swordfish bill is flattened in cross-section, anatomy of the swordfish rostrum. It is com­ lacks denticles in the adult (Carter 1919), and posed superficially of a thin layer of acellular usually contains a longitudinal series of central primary bone that has the appearance of dentine. chambers (Poplin 1975, 1976; Poplin et al 1976; The superficial layer is underlain by acellular Schultz 1987) that demonstrate considerable in­ vascular bone containing secondary osteons traspecific variation. In an X-ray examination (Haversian units) which are arranged into large of 39 rostra, I found 24 with chambers running bundles in the deeper regions (center) of the the entire length of the rostrum, five with cham­ sword. There are no comparable histological bers only in the distal one-half, seven with cham­ studies of istiophorid rostra. bers only in the distal one-fourth, and three with Lower jaw. With the exception of mentioning no chambers. There is a minor controversy over the presence or absence of the predentary bone, which bones form the swordfish rostrum. Gregory there are no published studies of other features and Conrad (1937) discussed the problem and of the lower jaw. The istiophorids have a stout concluded that the distal three-fourths is primarily mandible that is composed of the predentary, composed of premaxillae with paired nasals on dentary, and articular bones. There is a well­ the dorsal surface. The proximal one-fourth is developed intermandibular joint and each mandi­ composed of the dermethmoid, frontals, maxilla­ ble has a strong cone-shaped quadrate-articular ries, lateral ethmoids, and vomer. After dissecting joint. In contrast, the swordfish has a much a juvenile swordfish, Conrad (1937) concluded weaker lower jaw with no predentary bone or that the true nasal is "a minute but well-formed intermandibular joint. It has a rather simple bony element lying dorsad to the narial open­ quadrate-articular articulation. ings" and the bones that Gregory and Conrad Intrafamilial variation is unknown. The obser­ (1937) called nasals were just parts of the pre­ vation, however, that the shortbill spearfish has maxillae. To the best of my knowledge, all sub­ its predentary bone reduced to a tooth patch sequent workers (Nakamura 1983; Poplin et al (Fierstine and Applegate 1968), is incorrect 1976; Schultz 1987), except Johnson (1986), because the observation was based on an ab- have followed Gregory and Conrad (1937) and, not Conrad (1937). If most authors are correct, Table I. Relative length of denticulated area on dorsum ofrostrum offive species ofextant billfishes. Relative len[?th then X. gladius lacks the prenasal bones found was determined by dividin[? the length of the denticulated in istiophorids. However, if Conrad (1937) and zone by L (see Fig. 2 legend for definitio/l). Johnson (1986) are correct, then it is possible Number Average relative Range of that the bones they called parts of the premaxillae, studied length of relative and that others called nasals, are really pre­ Species (N) denticulated area length nasals. This matter needs to be resolved. Makaira nigricans 8 .02 .00-.03 In contrast, the istiophorid bill has paired pre­ Makaira indica 4 .05 .04-.07 nasal bones (Fig. I), is rounded in cross-section, Tetrapturus audax 6 .12 .03-.19 lacks any central chambers, and denticles adorn Istiophorus platypterus 4 .34 .08-.63 its ventral and generally its dorsal surfaces. The TetrapturuJalbiduJ 9 .75 .57·.78 BILLFISH PALEONTOLOGY 13

PM

M A

Figure I. Skull and lower jaw of a blue marlin (Makaira nigricans Lacepede, 1802). left lateral view. Composite illustration by Charles Bloomer primarily from a 43.3 kg (95.5 lb) '( caught off Kailua-Kona. HI. Bones labeled as follows: Articular (A). Dentary (D). Frontal (F), Lateral ethmoid (LE), Maxillary (M), Nasal (N), Predentary (PD). Premaxillary (PM), Prenasal (PN). 13

6 ...._-.1__...... __...1-__""-__...._....1 4 5 6 7 8 9 10

DEPTH X 10-1 .5l Figure 2. Simple regression of depth (D) and width (W) of 43 rostra measured at mid-rostrum length (.5L) for six species of billfishes. Rostrum length (L) was determined by measuring the straight-line distance from the distal tip to the anterior orbital margin of the lateral ethmoid. Istiophorus platypterus ('*'), n =4; Makaira indica ( X), n = 5; M. nigricans (e), n = 15; M. panamensis (0), n = I; Tetrapturus albidus (*), n = 9; T. audax (+), n =9. 14 PLANNING THE FUTURE OF BILLFISHES normal specimen. Additional specimens have a (Nakamura 1983). The surface of the scapula small, well-developed predentary bone and a that articulates with the large first pectoral ray lower jaw with an intermandibular joint. is flat in the black marlin, whereas the same Other skull bones. Billfishes differ in the surface in other istiophorids is round (Wapenaar proportions of their skulls. For example, the and Talbot 1964). Fierstine and Welton (1983) Xiphiidae have a short post-orbital region and noted that the articular surface of the first pectoral a longer rostrum, whereas the Istiophoridae have ray of the black marlin was about half the size a comparatively longer post-orbital region and ofthe same surface on a similar-sized blue marlin. shorter rostrum (Ovchinnikov 1970). In the The swordfish lacks pelvic fins and pelvic gir­ Istiophoridae, the cranium becomes stouter and dles (basipterygia), whereas istiophorids have wider with body length (Nakamura 1983). well-developed girdles with reduced number of Makaira has the widest and most massive rays. cranium and Istiophorus and Tetrapturus have Potthoff and Kelley (1982), Collette et aI more moderate-sized crania. (1984), and Potthoff et aI (1986) studied the early Vertebrae. Vertebral counts and shape differ osteological development of the median fin sup­ between the Xiphiidae and Istiophoridae, and ports in the swordfish and sailfish. Since these Makaira has enlarged transverse processes studies have few descriptions of adult structures, (apophyses) on its centra which are lacking in they have limited application to paleontology other istiophorid genera (Nakamura 1983). when most specimens consist of isolated rostral There are only a few other studies of vertebral fragments and vertebrae. morphology and variation. Gottfried (1982) noted that intervertebral Fossil Record foramina were positioned more anteriorly in Identification Makaira than in Istiophorus and Tetrapturus. Schultz (1987) proposed a classification scheme and that the posterior zygapophyses originated ofextant and extinct billfishes that differs signific­ more posteriorly on the centrum in Istiophorus antly from the scheme accepted by other modem than in the other istiophorids. Using the 14th workers (Collete et al 1984, Fierstine 1978, vertebra, Nakamura (1983) noted that Makaira Johnson 1986, Myer et al in press, Nakamura has wide and high (more or less square) neural 1983). His osteological observations were based spines, whereas Istiophorus has rather low, upon limited material: a skull preparation of a more narrow, triangular neural spines. Tetrapturus Mediterranean spearfish, and one rostrum each is more or less intermediate. The ratio of centrum of a blue marlin, sailfish, swordfish, and striped length to width of the 6th vertebra of a black marlin. Therefore, the distinguishing characters marlin, blue marlin, sailfish and striped marlin he proposed lacked an understanding of in­ differs among genera (Fierstine and Welton terspecific or intraspecific variation. Most of his 1988). The sailfish has the slimmest centrum, knowledge of the fossil specimens came from the black marlin and blue marlin the most robust, a thorough review of the literature, but not from and the striped marlin's centrum is intermediate actual specimens. Since I lack confidence in his in proportion. data, I can not accept his classification of The structure of the hypural complex differs taxonomic groups and lineages. between the Istiophioridae and Xiphiidae (Gregory Istiophoridae. Fierstine (l978) reviewed all and Conrad 1937; Fierstine and Walters 1968; fossil istiophorids known at the time. Most de­ Collette et al 1984; Potthoff and Kelley 1982; scriptions were based upon single isolated rostra Potthoff et al 1986; Johnson 1986). Similarities or vertebrae; all were described from Eocene, include three epurals, one uroneural, and a deep Miocene, Pliocene, and Pleistocene deposits, posterior notch between the upper and lower and few were compared with recent skeletal hypural plates. Differences include fusion of the material. Early workers (see references in parahypural with the hypural plate in istiophorids, Fierstine 1978 and Schultz 1987) generally whereas the parahypural is autogenous in Xiphias. placed specimens into new or existing fossil The uroneural remains free from the 5th hypural in species of Istiophorus, except for Van Beneden Xiphias while istiophorids lack a5th hypural element. (1871) who erected a new genus, Brachyrhynchus. Fins and fin supports. The black marlin is Although Fierstine (1978) questioned the accu­ the only istiophorid with a pectoral fin that can­ racy of these early identifications, he took the not be folded back against its body in the adult conservative approach and made no attempt to BILLFISH PALEONTOLOGY 15 re-identify or reclassify them. Myer et al (in press) believe that only M. The systematic studies of the recent Istio­ calvertensis is present in the very large Lee phoridae by Robins and deSylva (1960 and Creek billfish assemblage, but I think critical 1963), Nakamura et al (\968) and Nakamura review would probably synonymize M. calverten­ (\ 983) provided a firm basis and reference point sis with the blue marlin, and would find black for modem paleoichthyological investigations. marlin present as well (see paleozoogeographic Most fossil specimens are similar or identical discussion below). to recent specimens. Barbolani (1910) fore­ There are no known fossil specimens of the shadowed this interpretation when he placed genus Tetrapturus. Brachyrhynchus, plus some new specimens, Xiphiidae. Fossil swordfish are rare, with into the living /stiophorus herschelii (Gray, most specimens coming from the Italian 1838) which was synonymized with the blue Pliocene. Several rostral fragments that were marlin (Makaira nigricans Lacepecte, 1902) by originally named X. delfortrieri Lawley (\876), Nakamura et al (\968). were redescribed and renamed X. gladius var. The only fossil record of a sailfish is based delfortriei by Santucci (1923). Recently, Sorbini on a single, fragmentary vertebra from the Upper (\987) discovered a nearly complete 135-cm­ Pliocene San Diego Formation (Gottfried 1982). long swordfish in the Middle Pliocene Marecchia The fossil vertebra was compared to vertebrae River ichthyofauna. The specimen is figured but from only one or possibly two extant sailfish not described. In the only other positive fossil specimens, and was found similar. Although the record of the family, Poplin (1975) described a identification may be correct, it should be re­ rostral fragment of a swordfish from the subrecent studied by comparing it to a larger series of (Neolithic) of southern France. specimens from several billfish species. Other fossil records of Xiphias are erroneous The genus Makaira is represented in the or questionable. Xiphias rupeliensis Leriche, geological record from the Middle Miocene to 1908, was described from several vertebrae and present (Table 2). With the exception of the Lee an articular bone from the Middle Oligocene of Creek, Pungo River, and Italian specimens, each Belgium (Leriche 1908, 1926). The vertebrae discovery probably represents a single individual. are similar enough to Xiphiorhynchus kimbla­ Six species of Makaira have been recognized locki Fierstine and Applegate, 1974, that as fossils, the extant black marlin and blue marlin, Xiphias rupeliensis is probably a xiphiorhynchid and four extinct species, M. belgicus (Leriche, rather than a xiphiid. Eastman (\ 917) described 1926), M. calvertensis (Berry, 1917), M. courcelli a rather massive, isolated caudal vertebra from (Arambourg, 1927), and M. panamensis the Eocene of South Carolina as Xiphias (?) sp., Fierstine, 1978. Leriche (\926) described M. but it lacks one of the diagnostic features of an belgicus from a partial rostrum and abdominal xiphiid hypural, namely that the parahypural is vertebra, and Arambourg (\927) described M. not autonomous but, rather, is fused to the courcelli from a partial rostrum. These two hypural fan. Fierstine (1974) and Fierstine and species probably belong to the genus Makaira, Applegate (\ 974) thought they had found the but their incomplete condition makes it difficult rostrum of a new genus of Eocene swordfish, to confirm or reject their specific identification. but histological examination revealed that the

Table 2. Distribution offossil marlins of the genus Makaira.

Species Age Formation Locality References M. indica Early Pleistocene ?Cabatuan Luzon, Philippines Fierstine and Welton (1983) M. nigricans Pliocene .) Italy Barbaloni (1910) M. calvertensis Early Pliocene Yorktown Lee Creek, North Carolina, Meyer et al. (in press) U.S.A. Makairasp. Latest Miocene San Mateo Oceanside, California, Fierstine and Welton (1988) U.S.A. M. courcelli Latest Miocene Oran, Algeria Arambourg (1927) Makairasp. Late Miocene Monterey El Toro. California, Fierstine and Applegate U.S.A. (1968) M.panumensis Late Miocne Chagres S.S. Panama Fierstine (1978) M. belgicus Middle Miocene ? Belgium Leriche 1926) M. calvertensis Middle Miocene Pungo River Lee Creek, North Carolina, Meyer et al. (in press) U.S.A. 16 PLANNING THE FUTURE OF BILLFISHES rostrum was composed of calcified cartilage and based on a description of X. kimblalocki, that more properly belonged to a sawfish (Pristiidae) xiphiorhynchids are intermediate to the or saw shark (Pristiophoridae). Xiphiidae and Istiophoridae. The rostrum is A segment of an at1iculated vertebral column broad at its base, similar to a swordfish, but it with partial dorsal and anal fins, from the Middle is round in the distal one-half, similar to the Miocene of Poland, was identified as belonging istiophorids. Both the swordfish and Xiphior­ to a young specimen of Xiphias sp. (Jerzmanska hynchus have a central canal in their rostra, 1962). Comparing this account with Gregory while the istiophorids have only one pair of and Conrad (1937) and Potthoff and Kelley lateral canals. The abdominal vertebra of X. (1982), I believe that the vertebrae in the Polish kimblalocki is similar to the shape of an anterior specimen are too numerous and have the incorrect abdominal vertebra of the black marlin, whereas morphology to have anything to do withXiphias. the caudal vertebra is similar to the shape of a Xiphiorhynchidae. This monogeneric family swordfish caudal vertebra. (Xiphiorhynchus Van Beneden, 1871) contains five species [X. aegypticus Weiler, 1929, X. Paleozoogeography elegans Van Beneden, 1871, X. kimblalocki lstiophoridae and Xiphiidae. According to Fierstine and Applegate, 1974, X. parvus Eschmeyer et al (1983), Nakamura (1983), and Casier, 1966, and X. priscus (Agassiz, 1839)], Robins et al (1986), istiophorid billfishes are from the Eocene of either North Africa, western generally distributed in tropical and warm tem­ Europe, or eastern North America (see references perate seas, whereas the swordfish is distributed in Fierstine and Applegate 1974, Schultz 1987) world-wide in tropical (at mesopelagic depths), and one species (X. hungaricus Weiler, 1943) warm temperate, and cold temperate seas. Some from the Middle Oligocene of southcentral istiophorid species are found world-wide, while Europe (Weiler 1943). others have a more restricted distribution. For The rostra are round in cross-section, have example, the black marlin, shortbill spearfish, denticles (exact pattern is unknown), are com­ and striped marlin inhabit only the Pacific posed of acellular bone (Beltan 1976), and are Ocean, with occasional strays into the south­ distinguished from istiophorid bills by the eastern Atlantic Ocean, whereas the longbill presence of a large central canal and more than spearfish and white marlin are restricted to the one pair of lateral nutrient canals, and probably Atlantic Ocean, with the latter occasionally by the absence of prenasal bones. The original found in the Mediterranean Sea. descriptions Xiphiorhynchus were from cranial Fossil discoveries of the Istiophoridae and fragments that were presumed to have had rostra Xiphiidae are generally found in deposits within (Agassiz 1838). Woodward (1901), and most the distributional range of the extant species, recently Casier (1966), considered some of the and their discoveries have neither raised nor solved rostra and skulls conspecific, even though none any paleozoogeographic problems (Fierstine were articulated together. Until a xiphiorhynchid 1978; Fierstine and Welton 1983, 1988). skull is discovered with a rostrum, I lack confi­ Recently, I have been studying the rostrum dence in their identification. In fact, since most of a black marlin from the Middle Miocene of species are known only from rostral fragments, Panama. During the Middle Miocene there were I have little faith in the validity of the characters no known geographic or ecological barriers in that distinguish the species. the region of the future Isthmus of Panama to Schultz (1987) described a new genus and impede the movement of the black marlin from species, Thalattorhynchus austriacus, Middle the Pacific Ocean to the Atlantic Ocean (or vice Miocene, Austria, from a distal rostral fragment. versa). Gillette (1984) concluded there was a The distinguishing feature was a single, off­ Tertiary Caribbean Faunal Province that ex­ center, longitudinal canal that was observed by tended from Ecuador through the Caribbean Sea computerized tomography. No paired, lateral to coastal North Carolina in the Atlantic Ocean. nutrient canals were visible. The specimen looks Most likely, the black marlin was part of that similar to most istiophorid rostral fragments, faunal province and reexamination ofthe numer­ except for the single canal, which may be ous specimens of M. calvertensis from the Middle anomalous; therefore, I do not believe the speci­ Miocene deposits, Lee Creek, North Carolina men is a xiphiorhynchid. (Myer et al in press) will probably reveal that Fierstine and Applegate (1974) concluded, black marlin occurred in the Atlantic Ocean both BILLFISH PALEONTOLOGY 17

before and after the formation of the Panama true billfishes (Fierstine 1974) for the same land bridge. This early discovery weakens the reasons that Johnson found them a potential an­ hypothesis, "that the black marlin is a more cestor. Paleorhynchids have very elongated recent derivative (than the blue marlin) that was upper and lower jaws and numerous (45-60) unable to thrive in the Atlantic Ocean because vertebrae. of a temperature barrier" (Fierstine 1978), and After extensive analysis, Nakamura (1983) fuels the controversy that the black marlin may believes that the shortbill spearfish is most similar be more generally present, especially in the to the ancestral billfish. Some of the features south Atlantic Ocean, but lacks scientific confir­ are: bill short and round, cranium fairly stout mation. with a narrow parasphenoid and vomer, needle­ Xiphiorhynchidae. Xiphiorhynchids have like scales with few points, large slab-sided been found only in Tertiary deposits associated body, and triangular neural and haemal spines. with the north Atlantic Ocean and western His phylogenetic conclusions are weakened by Tethys Sea (or its remnant, the Mediterranean his mixing of plesiomorphic and apomorphic Sea). If xiphiorhynchids are the earliest known characters. true billfishes (see discussion below), then the No evidence has yet been presented to refute Atlantic Ocean and Tethys Sea are probably the the hypothesis that the Xiphiorhynchidae are center of origin for billfishes. morphologically intermediate to the Istiophoridae and Xiphiidae (Fierstine and Applegate 1974) Evolutionary Discussion and may be closest to the scombrid-billfish con­ Collette et al (1984) discussed the phylo­ nection postulated by Collette et al (1984) and genetic relationships of the six families and 44 Johnson (1986). genera of extant Scombroidei and concluded that the Istiophoridae and Xiphiidae are sister Summary groups, and together are a sister group to the The paleontology and relevant osteology of Scombridae. In a more extensive cladistic adult billfishes are reviewed from the literature analysis, Johnson (1986) concluded that the and unpublished studies. In the Istiophoridae, billfishes (Istiophoridae and Xiphias) should be the premaxillary and prenasal bones form a placed as separate tribes within the Scombridae, rounded rostrum with denticles, the lower jaw and together are a sister group to Acantho­ has a predentary bone and well-developed inter­ cybium. The last two groups share the feature mandibular joint, and the elongated vertebrae of a horizontal, premaxillary beak in the larvae, form a rigid spinal column. Certain skull and which elongates in adult billfishes. vertebral proportions exhibit intergeneric differ­ Although I do not believe that billfishes ences, and the denticle pattern and relative size belong to the same family as mackerels and of the nutrient canals of the rostrum show in­ tunas, I am now persuaded by the argument that terspecific differences. billfishes do not belong in their own separate All fossil istiophorids belong to one of five suborder (Xiphioidei) and are related to the species of Makaira. with the exception of one Scombridae. The similarities between billfishes vertebra specified as Istiophorus. There are no .and Acanthocybium is not a new idea and known fossil specimens of Tetrapturus. Johnson's novel presentation makes one look Istiophorids occur from the Middle Miocene to for evidence to support or reject his hypothesis. the present and have a zoogeographic distribu­ For example, does Acanthocybium have a pre­ tion similar to extant species, except that nasal bone? Does the istiophorid prenasal bone Makaira indica may have been common in the appear early or late in development? Does the western north Atlantic Ocean before the forma­ swordfish rostrum have prenasal bones or is it tion of the Isthmus of Panama. In the Xiphiidae, composed primarily of premaxillae? If scombrids the rostrum is flat, is composed of acellular bone are more primitive than billfish, then are there with secondary osteons, contains a variable fossil billfish having more than 24 vertebrae (or number of central chambers, and lacks denticles 26 in the swordfish) or fewer than three epurals? in the adult. There is a controversy over which Johnson (1986) believes that a thorough study bones form the rostrum. It may be composed of the extinct family Paleorhynchidae may resolve primarily of premaxillae, or premaxillae and some of these questions, but I doubt it. I con­ nasals, or premaxillae and prenasals. The lower cluded that paleorhynchids were probably not jaw lacks a predentary bone. The vertebrae are 18 PLANNING THE FUTURE OF BILLFISHES cube-shaped and are not modified into a stiffened Fierstine, H.L. 1978. A new marlin, Makaira panamensis. unit. from the Late Miocene of Panama. Copeia. 1978: I-II. Fierstine, H.L. and S.P. Applegate. 1968. Billfish remains Disregarding identifications based on isolated from southern California with remarks on the import­ vertebrae, the Xiphiidae is monotypic (Xiphias ance of the predentary bone. Southern Calif. Acad. gladius) and has no fossil record before Pliocene Sci. Bull. 67:29-39. times. The extinct Xiphiorhynchidae are charac­ Fierstine, H.L. and S.P. Applegate. 1974. Xiphiorhynchus terized by a rounded rostrum with central kimblalocki, a new billfish from the Eocene of Missis­ sippi with remarks on the systematics of xiphioid chambers and denticles, and apparently without fishes. Southern Calif. Acad. Sci. Bull. 73: 14-22. prenasal bones. The vertebrae are not formed Fierstine, H.L. and BJ. Welton. 1983. A black marlin, into a stiffened unit. The family is monogeneric Makaira indica. from the Early Pleistocene of the (Xiphiorhynchus) with six species, and is limited Philippines and the zoogeography of istiophorid billfishes. Bull. Mar. Sci. 33(3):718-728. to Tertiary deposits (Eocene of Africa, Europe, Fierstine, H.L. and BJ. Welton. 1988. A Late Miocene and North America, and the Oligocene of marlin, Makaira sp. (Perciformes, Osteichthyes) from Europe) associated with the north Atlantic San Diego County, California, U. S. A. Tertiary Res. Ocean and western Tethys Sea (or its remnant, 10(1):1-8. the Mediterranean Sea). The Xiphiorhynchidae Fierstine, H.L. and V. Walters. 1968. Studies in locomotion and anatomy of scombroid fishes. Mem. Southern may be closest to the scombrid-billfish connection Calif. Acad. Sci. 6: 1-31. postulated by recent workers. Gillette, D.O. 1984. A marine ichthyofauna from the Miocene of Panama, and the Tertiary Caribbean Faunal Province. J. Vert. Paleo. 4(2):172-186. Literature Cited Gottfried, M.D. 1982. A Pliocene sailfish lstiophorus Agassiz, L. 1838. Recherches sur les Poissons fossiles. platypterus (Shaw and Nodder, 1791) from southern Neuchatel. 5:89-92. California. J. Vert. Paleo. 2(2):151-153. Arambourg, C. 1927. Les Poissons fossiles d'Oran. Gregory, W.K. 1933. Fish skulls, a study of the evolution Materiaux pour la Carte geologique de I' Algerie. 1st of natural mechanisms. Trans. Am. Philos. Soc. Serie, Paleontologie. 6:1-298. 23(2):75-481. Barbolani, G. 1910. 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Poplin, C. 1976. Remarques sur I'analomie el "histologie Sorbini, L. 1987. Biogeography and climatology of Pliocene du rostrc de "Espadon (Xiphias g/adius L.). Rev. Trav. and Messinian fossil fish of Eastern-Central Italy . Boll. Inst. Peehes Marit. 40(3-4):716-718. Mus. Civ. St. Nat. Verona. 14 (1988): 1-85. Poplin, c., F. Poplin, and A. Rieqles. 1976. Quelques Van Beneden, P. J. 1871. Recherches sur quelques poissons partieularites analomiques et hislologiques du roslre fossiles de Belgique. Bull. Acad. R. Belg. CI. Sci. de "Espadon (Xiphia' g!adius L.). C. R. Seances 31 :493-518. Acad. Sci. Ser. D. 282:1105-1108. Wapenaar, M. L. and F. H. Talbot. 1964. Note on the Poplin, F. 1975. Resles de roslre d'Espadon trouves dans rigidity of the pectoral fin of Makaira indica (Cuvier). un gisement neolithique de I'etang de Leucate (Aude). Ann. S. Afr. Mus. 48:167-180. Bull. Soc. Prehist. Fr. 72:69-70. Weiler, W. 1943. Ein Schwertfisch-Rostrum aus dem Potthoff, T. and S. Kelley. 1982. Development of the ver­ Mittelologozan Ungarns. Z. Deut. Geol. Ges. 95:214­ tebral column. fins and fin supports, brachiostegal 217. rays, and squamation in the swordfish. Xiphias Woodward, A. S. 1901. Catalogue of the fossil fishes in gladius. Fish Bull. 80:161-186. the British Museum (Natural History). Vol. 4. 636 pp. Potthoff, T., S. Kelley, and J. C. Javech. 1986. Cartilage and bone development in scombroid fishes. Fish. Bull. Harry L. Fierstine received the B.S. degree 84:647-678. Robins, C. R. and D. P. de Sylva. 1960. Description and (biology) from California State University, relationships of the longbill spearfish, Tetrapturus be­ Long Beach, in 1957; the M.S. degree (ichthyo­ lone, based on western North Atlantic specimens. Bull. logy) in 1961, and the Ph.D. degree (zoology) Mar. Sci. 10:383-413. in 1965, both from the University of California Robins, C. R. and D. P. de Sylva. 1963. A new Western Atlantic spearfish, Tetrapturus pfluegeri. with a rede­ at Los Angeles. He has been a research associate scription of the Mediterranean spearfish. Tetrapturus in vertebrate paleontology at the Los Angeles belone. Bull. Mar. Sci, 13:84-122. County Museum of Natural History since 1967. Robins, C. R., G. C. Ray, and J. Douglass. 1986. A field He became a research affiliate to the Pacific guide to Atlantic Coast fishes of North America. The Gamefish Research Foundation, Kailua-Kona, Peterson Field Guide Series. Houghton Mifflin Com­ pany, Boston, Massachusetts. 354 pp. Hawaii, starting in 1987. He has published a Santucci, R. 1923. Nuovi studi sui pesci fossili della Liguria. number of papers on biIIfish paleontology, fish Boll. Soc. Geol. Ital. 41(3):195-213. anatomy, and ichthyology. He is currently as­ Schultz, O. 1987. Taxonomische Neugruppierung der Ober­ sociate dean for science and mathematics at the familie Xiphioidea (Pisces, Osteichthyes). Ann. Naturhist. Mus. Wien. Serie A. fUr Mineralogie and California Polytechnic State University, San Petrographie, Geologie und Palaontologie, Anthro­ Luis Obispo, California. pologie und Prahistories. 89:95-202.