0/ ISSN: 0098-4590 # Scientist '^Florida \J936j* Volume 67 Spring, Number 2004 CONTENTS Mass Occurrence of the Jellyfish Stomolophus meleagris and an Associated Spider Crab Libinia dubia, Eastern Florida Bjorn G Tunberg and Sherry A Reed Uptake of Phosphate and Nitrate Using Laboratory Cultures of Lemna 93 minor L Daniel P Smith, Matthew E McKenzie, Craig Bowe, and Dean F Martin Annotated Bibliography, 105 Donald R Richardson of Social Environment in Early Life on Cortical Depth, Locomotor Activity, and Spatial Learning in the Golden Mouse, 118 The Sand Pine Scrub Community: 1989-2001 An Effects Ochrotomys nuttalli Fred Punzo Report of Aplidium antillense Aplousobranchiata) from Florida First (Gravier, 1955), 144 (Tunicata, Thomas Stach 154 A Brief Description of the Courtship Display of Male Pike Killifish (Belonesox belizanus) LisaHorth Academy of Sciences Medalists Florida Endowment for the Sciences Florida 159 166 168 FLORIDA SCIENTIST Quarterly Journal of the Florida Academy of Sciences Copyright © by the Florida Academy of Sciences, Inc 2004 Co-Editor: Mrs Barbara B Martin Editor: Dr Dean F Martin Institute for Environmental Studies, Department of Chemistry, University of South Florida, 4202 East Fowler Avenue, Tampa, Florida 33620-5250 Phone: (813) 974-2374; e-mail: dmartin@chumal.cas.usf.edu Business Manager: Dr Richard L Turner Department of Biological Sciences, Florida Institute of Technology, 150 West University Boulevard, Melbourne, Florida 32901-6975 Phone: (321) 674-8196, e-mail: rturner@fit.edu http://www.floridaacademyofsciences.org The Florida Scientist is Inc., a non-profit scientific published quarterly by the Florida Academy of Sciences, and educational association Membership is open to in- dividuals or institutions interested in supporting science in plications may be its broadest sense Ap- obtained from the Executive Secretary Direct subscription is avail- able at $45.00 per calendar year or new interpretations of knowlof science as 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Dr John Trefry Department of Oceanography Florida Institute of Technology 150 W University Boulevard Melbourne, FL 32901 Past-President: Barry HDR Engineering, Wharton Gay Biery-Hamilton Rollins College 1000 Holt Ave., 2761 Winter Park, FL 32789-4499 Rebecca Amonett, Secretary e-mail: floridaacademyofsciences@osc.org Wharton Inc 2202 N Westshore Boulevard Suite 250 Tampa, FL 33607-5711 Secretary: Dr Elizabeth Program Chair: Dr Jeremy Montague Department of Natural and Health Sciences Barry University Miami Shores, FL 33161 Hays Barry University Miami Shores, FL 33161-6695 Published by The Florida Academy of Sciences, Inc Printing by Allen Press, Inc., Lawrence, Kansas Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Dean F Barbara Martin, Editor Volume 67 Spring, B Martin, Co- Edit or Number 2004 Biological Sciences MASS OCCURRENCE OF THE JELLYFISH STOMOLOPHUS MELEAGRIS AND AN ASSOCIATED SPIDER CRAB LIBINIA DUBIA, EASTERN FLORIDA Bjorn G Tunberg and Sherry A Reed Smithsonian Marine Station, 701 Seaway Drive, Fort Pierce, Florida 34949 Abstract: The jellyfish Stomolophus meleagris was collected both randomly and selectively in the Fort Pierce Inlet area of the Indian River Lagoon, eastern Florida on 26 and 28 March 2003 The total number ofS meleagris randomly sampled was 382 of which 16.5% carried an associated spider crab, Libinia dubia Two S meleagris carried two crabs each The male/female ratio of the crab was 0.82 The mean carapace width (CW) of the males was 22.9 mm and the females 20.0 mm The difference in size was significant between the sexes Crabs were only found on jellyfish with a 110 mm, while the bell diameter between 80 mm and range of the jellyfish was 70-130 mm More than twice as many females than males were found on jellyfish with a bell diameter of 80 mm, but otherwise the sex distribution was similar total size was no regardless of the size ofS meleagris There the relationship Key Words: between size (CW) and live significant difference between the sexes concerning wet weight Stomolophus meleagris, Libinia dubia, Indian River Lagoon, Florida Stomolophus meleagris is one of the most abundant species of scyphome- dusae along the southeastern and Gulf coasts of the United States (Mayer, 1910; Kraeuter and Setzler, 1975; Burke, 1976; Calder and Hester, 1978) According to Corrrington (1927) S meleagris is by far the most abundant scyphozoan of the South Carolina coast, and one of the more conspicuous planktonic organisms of the littoral habits zone The life history has been described by Calder (1982) and the feeding by Larson (1991) The spider crab Libinia dubia ocean waters and saltier estuaries is found on almost from nearshore The known range is from Cape Cod and Manning (1961) reported that to ca all 50 dubia 93 is m depth (Williams, 1984) Bahamas and Cuba Tabb common in Florida Bay and to southern Texas, L types of bottom in shallow 94 FLORIDA SCIENTIST [VOL 67 Dragovich and Kelly (1964) reported that it is the most common spider crab in Tampa Bay The larval development of L dubia has been described by Sandifer and Van Engel (1971) Jellyfish commonly harbor commensal forms, with certain symbionts being characteristic tive For instance, some brachyuran crabs exhibit protective and transpor- forms of commensalism with jellyfish S meleagris and L dubia was between L dubia and also describes the association information on this association been reported In an is The association between S meleagris However, the very limited and no detailed observations have earlier study, between Cancer gracilis and (Trott, 1972) reported by Corrington (1927) Williams (1984) first Weymouth (1910) described the association jellyfish (species not determined), and Trott (1972) described the relationship between Stomolophus nomurai and the portunid crab Charybdis feriatus Young blue crabs, Callinectes sapidus, have frequently been observed clinging to the umbrellas of the sea Chrysaora quinquecirrha nettle, (Jachowski, 1963) A mass occurrence of S was observed surface, that some with meleagris, body March 2003 crabs clearly visible on the in the Fort Pierce inlet area, eastern Florida in allowed analysis of the association Methods —Random sampling of the S meleagris was performed on 26 and 28 March 2003 from Lagoon (IRL) by using fine mesh dip nets The bell diameter was cm, and each specimen was thoroughly examined for L a small boat in the Fort Pierce Inlet area of the Indian River (Figure Each ) measured with a dubia If a crab jellyfish was transferred to a container filled with saltwater plastic ruler to the nearest was found, were then transferred it was measured (carapace width) 28 March, selective sampling was also performed, them were collected the laboratory and the bell The to the nearest mm and sexed All crabs to another container filled with saltwater for further treatment in the laboratory On 31 March, relationship all i.e., crabs sampled on 28 between the bell On only jellyfish with visible crabs associated with March were weighed to the nearest 0.1 g in diameter of the jellyfish and the crab carapace width diameter and the sex distribution of the crabs were calculated from the random sampling data Results —Twenty-eight males, 34 females and juvenile, of L dubia were found when random specimen, presumably a collections of S meleagris (382 specimens) were performed on 26 March, which corresponds to a male/female ratio of 0.82, and an association rate of 16.5% The size distribution of L dubia of the randomly collected S meleagris of the males was 22.9 is presented in Figure The mean carapace width mm (SD = 4.8 mm) and the females 20.0 mm (SD = 4.0 mm) The size difference was significant between the sexes (p = 0.004, Mann Whitney Rank Sum test) The crab size distribution of the selectively collected jellyfish is presented in Figure The mean carapace width of the males was 23.2 mm (SD = 4.8 mm) and the females 23.5 mm (SD = 4.0 mm) The male/female ratio of the crab on the selectively collected jellyfish was 2.14 (62 males and 29 females) The size difference was not significant between the sexes (p = 0.72, unpaired t-test) The selectively sampled females were significantly larger (CW) than the randomly sampled ones (p = 0.003, significant size difference Mann Whitney Rank Sum between the males (p = 0.561, test), but there was no unpaired t-test) TUNBERG AND REED—JELLYFISH AND SPIDER CRABS No 2004] 95 27°29'N 80°19'W Fig The Fort Pierce of the Indian River Lagoon, eastern Florida Sampling of Inlet area Stomolophus meleagris was performed on 26 and 28 March 2003 within SMS = the area marked with a rectangle Smithsonian Marine Station The male/female ratio of all crabs combined was 1.43 (90 males and 63 females) During the selective sampling on 28 March two specimens of S meleagris had two crabs attached, one 19 (bell diam = 100 mm) the other (bell diam = 90 mm) The mm) (CW = 30 mm and (CW = 28 mm and 19 mm) with two males with two females relationship between carapace width and wet weight for males and females was achieved by using polynomial regressions Both regressions were highly significant: p < 0.001 (power of performed test with alpha = 0.050: 1.000) There was no significant difference in size or weight between is presented in Figure the females and males (p Sum The best fit = 0.1 14 and p = 0.081, respectively) (Mann-Whitney Rank Test) The and the relationship bell nificant correlation females (p The between the carapace width of males and females of L dubia diameter of S meleagris between = 0.1 15) bell is presented in Figure There was no sig- diameter of the jellyfish and the carapace width for and males (p = 0.469) (Pearson Product Moment Correlation) between bell diameter of the jellyfish and the sex distribution relationship of the crabs is presented in Figure More than twice as many females than males 96 FLORIDA SCIENTIST 10 15 [VOL 67 20 25 30 35 CARAPACE WIDTH (mm) Fig Size distribution of Libinia dubia from Stomolophus meleagris collected randomly on 26 and 28 March 2003 were found on jellyfish with a bell diameter of 80 mm, but otherwise the sex was similar regardless of the size of S meleagris Figure shows the number of crabs found on different size classes of S meleagris As presented earlier, crabs were only found on jellyfish within size distribution TUNBERG AND REED—JELLYFISH AND SPIDER CRABS No 2004] 10 20 15 25 30 97 35 40 CARAPACE WIDTH (mm) Fig Size distribution of Libinia dubia from Stomolophus meleagris collected selectively on 28 March 2003 classes 80-1 10 mm jellyfish mm The highest percentage of crab occurrence was recorded on 80 (22.6%) and the lowest (10.9%) on those measuring 110 Discussion were abundant —Mayer (1910) in mm reported that mature individuals of S meleagris winter and spring off the coast from Florida to South Carolina FLORIDA SCIENTIST 98 [VOL 67 25 fern width vs fern weight females male width vs male weight plot males plot Females 20 Coefficients: b[0] 12.23 b[1] -1.75 0.09 b[2] b[3] \- r I CD ED - 9.01 e-4 0.981 Males 15 Coefficients: b[0] -23.02 b[1] LU b[2] b[3] r LU 3.12 -0.14 2.38 e-3 0.977 > 10 - 10 15 20 25 30 35 40 CARAPACE WIDTH (mm) Fig relationship between carapace width (CW) and live wet weight of Libinia dubia and was mostly confined to ocean water off However, Kraeuter and Setzler (1975) performed studies on S meleagris in Georgia estuaries and concluded that this species does occur offshore in the winter, but that it also spends much of its early life in sounds and estuaries Large individuals appear offshore in March and apparently move in nearshore in May and It was The rarely seen in brackish harbors, the coast June Small individuals were collected in early July in the estuary These populations TUNBERG AND REED—JELLYFISH AND SPIDER CRABS No 2004] 99 /LO D 26 - 24 - # vs females bell diam bell diam vs males • 22 o - • • Q E E I H Q 20 O - • • 18 - 16 - • • LU O < CL < < o • • 14 - • 12 - 10 - « 80 90 110 100 BELL DIAMETER (mm) Fig The relationship between bell diameter of Stomolophus meleagris and carapace width of males and females of Libinia dubia decreased in numbers by August After mid-October until March According and October 1987, from June to to Larson (1991) throughout the year in all individuals disappeared performed between June 1986 Gulf of Mexico, S meleagris was abundant Burke (1976) reported that S meleagris was found Mississippi Sound, with the highest abundance during in the north-eastern October in a study FLORIDA SCIENTIST 100 [VOL 67 CO DO < a: o LU CD 90 100 BELL DIAMETER (mm) Fig The relationship between the bell diameter of Stomolophus meleagris and the sex distribution of Libinia dubia midwinter Specimens collected ranged in size from to 380 suggesting that a few of these medusae may mm bell diameter, survive for longer than a year is usually most abundant during late Even though we observed a few jellyfish far from In the Fort Pierce Inlet area, S meleagris summer and early fall (pers obs.) the inlet in the IRL, the highest abundances were near the Fort Pierce Inlet FLORIDA SCIENTIST 156 Fig Line drawings of a single entire zooid, a larva, and a cross section through the stomach region (plane see arrow) of Aplidium antillense as in — — — —rudiment of stomach, — rst [VOL 67 intestine, la languet, larva, lg sc lm — — atrial siphon, en — endostyle, fp longitudinal muscle, lp statocyte complex, sd —sperm — — fecal pellet, — — larval papilla, oc duct, st —stomach, ta —gonoduct, ov — — gd ocellus, tail, te ovary, testes STACH— NOVEL ASCIDIAN No 2004] 157 Bermudas Fort Pierce, Florida w Kingston, ©Guadeloupe Jamaica Fig Map showing localities of earlier (circles) and present (arrow) findings of Aplidium antillense The zooids of A or even orange was f Martinique exile are variably colored as described The living zooids of A antillense by Van brightly colored, yellow to orange Colonies of A exile Discussion —Aplidium is Name (1945), sometimes brown, yellow, from Florida were of a dull gray color 1902), A exile 1898) from Tortugas his doctoral thesis, Bingham Lagoon (Bingham, United States so Name (1945) pellucidum (Leidy, 1855), A constellatum mudae (Van Name, (Sluiter, from many observers Van attract attention (Verrill, 1871), A (Van Name, 1902) from A stellatum lists (Verrill, 1871), A ber- Florida, and A funginum appended to 1990) A antillense has not been reported from the continental far Several hypotheses could account for this misidentification might be reasons Van Name in his original description of A exile mentions that the species possesses (Van Name, 1902) (Van Name, 1945) species A exile and A constellatum from the Indian River Taxonomic inconsistency and/or folds stomach their small size Island In an informal identification key, lists the 200 described a genus that contains about Although they are generally clearly distinguished from one another, does not Only from the same location were brown It is In his major work, he corrects this number to stomach 20 or more conceivable that he was dealing with two different species Gravier (1955) distinguished A antillense from A exile by: a smaller number of rows of stigmata (never more than 12) plications (12-14), the subdivision in A antillense, a of the smaller intestine into number of stomach three parts, and a comparatively longer postabdomen Monniot, on the other hand realized that FLORIDA SCIENTIST 158 [VOL 67 most of these characters were too variable and overlapping to be reliable for species acknowledged only the number of stomach A antillense versus more than 20 and up to 35 in A exile, identification This author 11-14 in distinguishing characteristic to identify A antillense (Monniot, 1983) On this ground the latter clearly distinguishes A antillense 1-13 in the Floridian from A The number of stomach exile specimens of A antillense, whereas is fact that A antillense in the present study might be a reason facilities Caribbean traffic in the slightly to ships' hulls or in the bilge Finally, it last 48 years since Acknowledgments This is at — I Because ship might have been transported attached that the its number of reported observations renders Marine Station in the vicinity of water of ships might be also conceivable changed over the exile was found only to consider another hypothesis intense, A antillense is It is number is longer compared to its in A exile this diameter in A antillense versus a more globular shape in A harbor plications also from the Indian River Lagoon always higher than 20 In addition, the stomach The as a valid 1972; Monniot, author could discern between the two similar species co-occurring in the same geographical areas plications, geographic range of the species However, the low original description hypothesis hard to this test thank the former director Dr Mary E Rice and the staff of the Smithsonian Reed Fort Pierce for support during this study, especially S for guidance in the field Smithsonian Marine Station Contribution No 573 LITERATURE CITED Bingham, B L 1990 The ecology of epifaunal communities on prop roots of the red mangrove, Rhizophora mangle Ph.D Goodbody, I dissert Florida State Univ Tallahassee, FL 2000 Diversity and distribution of ascidians (Tunicata) in the Pelican Cays, Belize Atoll Res Bull 480:304-326 2003 University of the West Indies, Jamaica, Pers Gravier, R 1955 Ascidies recoltees par Rev Trav Leidy, Inst, scient Commun "President Theodore Tissier" (Campagne de printemps New Monniot, Jersey F 1972 J 1951) Peches mark 19:611-631 1855 Contributions towards a knowledge of the marine fauna of the coasts of J Acad Nat Sci Philadelphia 2(3): Rhode Island and 135-152 Ascidies aplousobranches des Bermudes Polyclinidae Hist, nat., Paris 3e ser (no le et Polycitoridae Bull Mus nat 82):949-962 1983 Ascidies littorales de Guadeloupe III Polyclinidae Bull Mus nat Hist, nat., Paris 5(2):413^122 Sluiter, C P 1898 Tuniciers recueilli en 1896, par la Chazalie, dans la mer des Antilles Mem Soc Zool France 11:5-34 Van Name, W G 1902 The ascidians of the Bermuda islands Trans Connecticut Acad Sci 11:325- 412, plates 46-64 1945 The North and South American Ascidians Verrill, A E 1871 Descriptions of some imperfectly Amer J Sci 3(1):54^146 Florida Scient 67(2): 154-158 2004 Accepted: September 23, 2003 Bull Am Mus known and new Nat Hist 84:1-475, 31 plates ascidians from New England Biological Sciences A BRIEF DESCRIPTION OF THE COURTSHIP DISPLAY OF MALE PIKE KILLIFISH (BELONESOX BELIZANUS) Lisa Horth Kellogg Biological Station, Michigan State University, 3700 East Gull Lake Drive, MI 49060 Hickory Corners, Abstract: The mating behavior of the different live-bearing fish species within the family Poeciliidae varies widely Several species, such as guppies (Poecilia reticulata), sailfin mollies (Poecilia and swordtails (Xiphophorus latipinnaj helleri) are well-studied However, very been recorded regarding the largest poeciliid, the pike killifish little information has (Belonesox belizanusj This work documents the courtship behavior of male pike killifish, a species which persists within the United States, only in Florida Unlike males in the non-courting Poeciliidae species, pike killifish males repeatedly conduct particular behavioral acts when A behaviors in the presence offemales Females not conduct courting male will position himself in front and gonopodium (male mating becomes more vibrant during a of, and perpendicular to which is of His coloration structure), in her direction, prior to attempting to mate display, this suite a female, and fan his fins often followed by mating attempts Pike killifish compose a monotypic genus that has previously been used as an outgroup when studying relationships between live-bearing species Thus, documenting the courtship behavior of this species studies is where mating-behavior comparisons are being made across genera within Key Words: Mating behavior, live-bearing fish, teleost, Understanding the mating behavior of closely information for testing selection behavior Panhuis many sorts of hypotheses, Poeciliidae related taxa provides useful such as those involving sexual in which male mating Uy and Borgia, 2000; and speciation Such studies often draw comparisons is of prime importance (Seehausen et et al., relevant for use in this family al., 1997; 2001; Boughman, 2001) Mating behavior data have also been used in cladogram construction Cladograms are then assessed relationships identified in As an example, such them reflect to determine whether the molecular phylogenies (Paterson cladistic analyses have been used to et al., 1995) how determine well phylogenies account for social organization in primates (Di Fiore and Rendall, 1994) These types of comparisons are relevant in the live-bearing fishes, as well In swordtails {Xiphophorus helleri), there exists a standing hypothesis that females prefer to mate with males possessing an exaggerated male elongated caudal 1990a, b) fin), and This hypothesis was recently called into question Present address: Section of Evol CA trait, the sword (an that this preference arose in swordless species (Basolo, And Ecol., Storer Bldg., 95616-5270 159 when character states Shields Ave., University of California at Davis, Davis, for [VOL.67 FLORIDA SCIENTIST 160 male tail-traits were mapped onto molecular phylogenies (Meyer et al., 1994; see Marcus and McCune (1999) used phenotypic and DNA sequence construct data to a Xiphophorus phylogeny They found a correlation between increased body size and sword growth rate, and suggest increases in these traits also Meyer, 1997) environmental relates to high visibility for females by phylogeny- wide female preference Their hypothesis for the presence of a strengthened is sword (Basolo, 1995) Marcus and McCune (1999) additionally suggest that selection for small body size in male X continens is derived, and may contribute to the loss of the sword despite the presence of female preference for a sword in this s wordless species The mating behavior of many of the fishes that compose the live-bearing family, Poeciliidae, has been well studied (Farr, 1989) Though a complete phylogeny of this family remains to be published, smaller phylogenies of groups within the family have been constructed (Meyer et al., 1994; Morris et al., 2001) Such phylogenies are sometimes created using the pike killifish as an outgroup (Lydeard et al., 1995a, b) However, very little mating behavior data has been gathered for this monotypic genus In general, mating behavior in the family varies markedly from species to species Some species exhibit conspicuous male displays (e.g., sailfin mollies, Woodward, Poecilia latipinna, Travis and 1989), and female preferences guppies, Poecilia reticulata, Houde, 1988), while others for such behaviors (e.g Limia sp., Farr, reticulata) there exists a correlation little to (e.g., no evidence 1984) For example, in guppies {Poecilia between female preference and males with bright color-spot patterns (Houde, 1987; is show Houde and Endler, 1990) However, there also a higher cost of predation for males that display bright spots (Endler, 1980) Within populations of sailfin mollies {Poecilia latipinna), females prefer larger- bodied males, which have larger dorsal fins (that are typically higher rates of display (Ptacek occur among used in courtship displays), populations in size-specific male courtship rates (Ptacek and Travis, 1996) In contrast, gila topminnow {Poecilipsis occidentalis) males conduct very simple courtships displays (Constanz, 1975) Finally, least formosa) males express no obvious color or size killifish {Heterandria polymorphisms and are not known females (Rosen and Tucker, 1961) to display to The pike poeciliids and and Travis, 1997) Large differences also killifish {Belonesox belizanus) is the largest-bodied and forms a monotypic genus within the tribe (—105 mm) of the Gambusiini, in the subfamily Poeciliinae (Meffe and Snelson, 1989) This piscivorous species is native Mexico and Central America (Rosen and Bailey, 1963) and was introduced to Dade County, Florida in 1957 (Belshe, 1961), where it has remained rare, but persisted Currently, extant pike killifish populations in Dade County are found in a small number of freshwater ditches The species has not expanded its range outside of south Florida The limited descriptor of the species' reproductive behavior to includes that used in a broad comparison of the poeciliids (Rosen and Tucker, 1961) Rosen and Tucker (1961) simply indicate that the species has a mating display, including the demonstration of a 'specific orientation' by the males, and a lack of mouth/body contact to describe the However, it was not relevant to the mating behavior more thoroughly message of that publication A second account states that males demonstrate a simple courtship display only after females give birth (Farr, 1989) HORTH— PIKE No 2004] KILLIFISH COURTSHIP DISPLAY Farr's reasonable rationale regarding reduced display periods whom wait' predators for 161 is that these are may a high level of sexual activity 'sit and deter prey Belshe (1961) also mentions an extended period of copulatory attempts, but does not describe mating behavior any further The objective of the present information about the courtship display of pike work is to clarify from a field collected killifish population in south Florida Methods from canals in — Florida Though the fish are distinctive water surface), they are quick and often found each fish that was MI) and housed by m received collected in a August 1995, the wide by full sun and two females) were collected via cast-net In June 1995, fifteen pike killifish (13 males Dade County, fish The m were moved deep to 380-L the water and adorned with clear, rectangular Plexiglas The aquarium was located Sand was placed on long, slender, swim near the Kellogg Biological Station (Hickory Corners, plastic vat, filled with well to a (—3-6" and around vegetation, so several casts were made for were transported fish 1-KL dark-gray in m directly beneath a bottom of the aquarium, and artificial plants aquarium, measuring artificial by m 1.5 In m long window and plants were anchored in it Live guppies (Poecilia reticulata) were added daily, as prey During autumn, heaters were used to maintain a 22°C water temperature Observations were and evening for made at least daily, from July to October, on the Fish were observed in the morning fish 30 minutes, and were checked on several additional times during the day to determine whether displays were occurring throughout the day Data were recorded daily regarding the sex of the fish observed, whether or not the fish displayed, and what behaviors composed the display Video recordings were made of the mating behavior and display, as The mating well comprised, in part, of several behaviors previously described in other poeciliid species Kadow, 1954; caudal fin spread: Baerends et al., (e.g., ritual was intromission: 1955; post-copulatory jerking: Liley, 1966; Parzefall, 1969, 1979; Farr, 1977) Results and Discussion —The activity of the pike killifish when located in the gray vats included simple behaviors: rapid darting from behind plants to capture prey, rapid and swimming from behind one in close proximity plant to another, and occasional typically dark-gray, occasionally light gray In contrast, within a swimming parallel, another individual The body-color of fish in the vat was to, No other activity few days of being transferred was observed to the transparent Plexiglas housing, males were seen displaying to females Females did not display Both sexes appeared brighter in color in the clear aquarium, their body-colors being silvery-gray to white abdominally, with the presence of golden hues dorsally from dark (e.g., to light gray in lower vertebrates in the dispersion of melanin, a black In general, morning male displays occur until dusk persist, as in dermal cells change in hue from a change (Hadley, 1996) several times a day, from early Displays occurred daily during the entire study period of June through October Displays may pigment present intermittently, A frogs) often results last anywhere from a few seconds to many minutes, and repeated acts, for the majority of a day Often, a male displays for several minutes, then leaves the female and 30 sec), then returns to the female, swims around and continues for a short period (e.g., to display for several more minutes This was a very typical observation All males displayed to females frequently During a mating display, the central portion of a male's body, inclusive of the gonopodium (modified fin used for mating), is near the female's nasal The male maintains his position in front of the female while the central portion of his body from side-to-side in a sigmoid-motion dorsal fin and and oral cavities moving just FLORIDA SCIENTIST 162 He swings his gonopodium perpendicular (a modified anal of his long-axis the to used to inseminate the female) fin body and toward simultaneously rapidly fanning his erect dorsal Generally, poeciliid males swing the [VOL 67 female, the while the direction of her snout fin, in gonopodium forward and backward in a more The traditional movement does linear motion, in preparation for, or during mating not include distinct sideways thrusts, as occurs during these displays During a mating-display, a female typically remains motionless and maintains her position in clear view of the male Sometimes a female will shift her position and change the angle between herself and the displaying male The male then repositions himself in front of the female, and resumes fanning in her direction female swims away just prior her and continues to display male may to, On occasion, the or during the-display Typically the male follows When another female swims near the pair, the courting re-direct his display to the new female After fin fanning, the male sometimes positions himself beneath, and parallel to, the female, his snout near her gonopore topminnow (Poecilipsis occidentalis) Molly (Poecilia latipinna) and gila males conduct gonoporal nibbles of the female urogenital pore (Travis and Woodward, 1989; Fair, 1997; Constanz, 1975) when in this position urogenital pore, I which did not see any nibbles or any male contact with a female's Rosen and Tucker mouth wide, arc gonopodium forward, then backward slowly, and in consistent with the observations of is (1961) While beneath a female, a male will occasionally open his his body slightly, and swing his the traditional pattern of live-bearing males (parallel, not perpendicular to the axis of his body) The gonopodial swing and mouth-opening behavior that display for a long period, but that also occurs in males have not moved beneath the female After displaying, a male occasionally attempts to copulate, and swings his gonopodium directly toward the female's gonopore, other poecillids (more parallel than perpendicular to his attempts to intrude while a female the potential new suitor intruder and the female is own manner consistent with body) If a second male being courted, the courting male either chases away, or jockeys to maintain his position between the The defending male swims to her, holding his position in a parallel to the female, very near between the female and the intruding male During these encounters, and sometimes independent of a female, the two males will parallel to one another, displaying their throat-patches at Additional changes in the male's appearance when courting a female include a deepening in hue of the typical gray to light-gold body-coloration, to a deeper gold A small pelvic fin, swim one another much near the gonopodium, turns deep gold and pink In body regions also turn pink: 1) a small area on the head, just posterior an area on the body immediately ventral to the dorsal fin, and 3) the area on the body anterior to the tail fin The caudal fin, though typically translucent, displays a black, translucent hue during courtship The caudal fin-spot (present in all individuals in this collection, but not visible at all times), and horizontal rows of black, body-spots, darken How the body-spots and caudal finspot display is used in these, and other poeciliids (e.g Heterandria Formosa, Poecilia latipinna), remains to be determined Pike killifish males also have two pouches of skin, one under the mouth on the throat, and one on the upper abdomen, addition, three to the otoliths, 2) HORTH—PIKE No 2004] KILLIFISH COURTSHIP DISPLAY which they sometimes distend during a display 63 to a female, or in an aggressive encounter with another male In contrast to males, females not display Their primary mating-behavioral repertoire appears to consist of either remaining motionless while a nearby male moving away from a displaying male Females occasionally open their mouths wide when being courted, potentially allowing them to receive chemical cues from a fanning male However, the actual purpose of this mouth-opening behavior is not known Females gave birth at least three times during the study period On all occasions, the young were cannibalized within two days Males courted displays, or females both before and after parturition In the collection of 13 males and females that mating-displays toward females I made only one I had, all males conducted observation out of hundreds, thrust his gonopodium at a female without Lack of a display is consistent with Fair's observations (1989) In addition, if the larger males were displaying to the females, since there was a malebiased sex ratio, the smaller males sometimes displayed to other males, though when they had the opportunity they would preferentially display to females Fair (1997) where a male turned upside-down and displaying first has noted that in guppies (Poecilia reticulata), when the sex ratio is male-biased, males will display to other males Gambusia holbrooki males also occasionally attempt to mate with other males (pers obs.) With respect killifish, it may because he was not using the equivalent of a to intense sunlight, as was used a subset of the pattern that more activity, noted in other studies of pike to the lack of mating-displays be that Fair (1989) did not observe the mating-display that I clear, oversized-housing, did I exposed in this study Indeed, his behavioral descriptor is observed Even in my study, there appeared to be much both with respect to behavior and color-change, in the brightly sunlit Plexiglas chamber, than in the dark gray vat was not designed However, this was observational study as an empirical manipulation so confirmation of the light to the display is not possible from this importance of work Observations on these fish in nature would be most helpful for understanding the significance and frequency of well as that documented here, male displays may prove ing mating behavior in poeciliids, or to females Such information, as useful to those developing theories regard- mapping characters such as mating behavior onto phylogenies to identify the role of phylogenetic constraints on behavioral development Acknowledgments —Thanks unpublished observations, supplies, W R to Cailteaux, J Trexler for comments, Loftus and A Turner for assistance in collecting J fish, Farr for sharing G Mittelbach for and C Huckins for the hand-made Plexiglas aquarium This work was supported by Research Training Grant DBI 9602252 MSU AUF NSF 04/94-106-01 LITERATURE CITED Baerends, G P., (Peters) I R Brouwer and H An analysis of the T Waterbolk 1955 Ethological male courtship pattern studies Basolo, A L 1990a Female preference predates the evolution of the sword Washington 250:1426-1427 on Leibistes reticulates Behaviour 8:249-335 in swordtail fish Science FLORIDA SCIENTIST 164 [VOL 67 1990b Female preference for male sword length in the green swordtail, Xiphophorus helleri (Pisces: Poecillidae) Anim Behav 40:332-338 1995 Phylogenetic evidence for the role of a pre-existing bias in 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physiology in the guppy, Lebistes reticulatus (Peters) Ph D Dissertation Liley, N R New York Univ New York, NY 1966 Ethological isolating mechanisms in four sympatric species of poeciliid fishes Behaviour, Suppl 13:1-197 Lydeard, C M., C Wooten, and A Meyer 1995a Molecules, morphology, and area cladograms: a cladistic and biogeographic analysis of Gambusia (Teleostei: Poeciliidae) Syst Biol 44: 221-236 1995b Cytochrome b sequence variation and a molecular phylogeny of the live-bearing fish genus Gambusia (Cyprinodontiformes: Poeciliidae) Can Marcus, J M and A R McCune J Zool 73:213-227 1999 Ontogeny and phylogeny in the northern swordtail clade of Xiphophorus Syst Biol 48:491-522 Meffe, G K and F F Snelson 1989 Ecology and Evolution of Livebearing Fishes (Poeciliidae) Englewood Cliffs, NJ Meyer, A 1997 The evolution of sexually Prentice Hall, Poeciliidae) Heredity , J Morrissey, and M Shartl Xiphophorus selected traits in male swordtail fishes {Xiphophorus: 79:329-337 fishes inferred 1994 Recurrent evolution of a sexually selected trait in from a molecular phylogeny Nature 368:539-542 Morris, M R., K DeQueiroz, and D C Morizot 2001 Phylogenetic relationships among populations of northern swordtails {Xiphophorus) as inferred from allozyme data Copeia 2001:65-85 Panhuis, T M., R Butlin, M Zuk, and T Tregenza 2001 Sexual selection and speciation Trends Ecol Evol 16:364-371 Parzefall, J 1969 Zur vergeichenden Ethologie verschiedener Mollynesia-Arten einschliesslich einer Hohlen von M Sphenops Behaviour 33:1-37 HORTH—PIKE No 2004] 1979 KILLIFISH COURTSHIP DISPLAY 65 Zur genetic und biologischen Bedeutung des Aggressionsueshaltens von Poecilia sphenops (Pisces, Poeciliidae) Z Tierpsychol 50:399-422 Paterson, A M., G P Wallis, and R D Gray 1995 Penguins, petrels and parsimony: does cladistic analysis of behavior reflect seabird phylogeny? 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