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Cirripedia: Thoracica) Author(S): Kristina M Comparison of Larval and Adult Stages of Chthamalus dalli and Chthamalus fissus (Cirripedia: Thoracica) Author(s): Kristina M. Miller, Sally M. Blower, Dennis Hedgecock, Jonathan Roughgarden Source: Journal of Crustacean Biology, Vol. 9, No. 2 (May, 1989), pp. 242-256 Published by: The Crustacean Society Stable URL: http://www.jstor.org/stable/1548504 Accessed: 16/02/2009 12:15 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=crustsoc. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. The Crustacean Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Crustacean Biology. http://www.jstor.org JOURNALOF CRUSTACEANBIOLOGY, 9(2): 242-256, 1989 COMPARISON OF LARVAL AND ADULT STAGES OF CHTHAMALUS DALLI AND CHTHAMALUS FISSUS (CIRRIPEDIA: THORACICA) Kristina M. Miller, Sally M. Blower, Dennis Hedgecock, and Jonathan Roughgarden ABSTRACT Adult and larval forms of Chthamalusdalli and Chthamalusfissusare comparedand distin- guishingcharacteristics described. Adult forms of these species differin malate dehydrogenase enzyme allozymes, in the setae of the second cirrus,and in shell characteristics.Larval forms are morphologicallyidentical, although larvae culturedfrom adults collectedat MontereyBay differin size at naupliarstages I-III and at the cyprid stage. Chthamalus dalli Pilsbry is a common the MDH allozymes, by the setae of the northern barnacle inhabiting the high in- second cirrus (W.A. Newman, personal tertidalzone from Alaska to San Diego and communication), and by a combination of northernJapan. ChthamalusfissusDarwin, externalfeatures. The larvaeof these species a close relative of C. dalli, is a southern appear to be morphologicallyidentical. barnacleinhabiting the high intertidalzone MATERIALSAND METHODS from San Francisco to Baja California. In the between San Francisco and San Mass CulturingTechniques.- Ripe eggswere obtained region from freshlycollected adult barnaclesand placed in 1 Luis Obispo, these species areabundant and um-filteredsea water.Newly hatchedlarvae were col- can be found in combined densities reach- lected by pipettingthem from areasof maximumlight ing 70,000 per m2in the same high intertidal intensity created by a point light source. The larvae band on the shore Newman and Abbott, were then placed in a 4-1 glass jar at a density of one (cf. larva per 2 ml sea water.The jars, each of which con- 1980). tainedfresh 1 ,m-filtered sea waterat 33.5 ppt salinity, The community structure of barnacle werekept at either 13?or 18?Cand mildly aeratedwith populationshas been studied extensively in filteredair from a 1/3hp oil-freepump. The larvaewere the studies fo- fed the Tahitianstrain of Isochrysisgalbanaat a density past twenty-fiveyears. Early x cused on the roles of and of 1 105cells per ml. competition pre- The waterin the jars was changedon alternatedays dation on adult barnacles in structuring by pouringthe contents through100-gm filters,wash- benthic communities (Connell, 1961a, b; ing the larvae in the filters (continuallyimmersed in Southward, 1976). Recent work has ex- water)with 1 gm-filteredsea water,placing the larvae the role of settlementin benthiccom- in small bowls, and pipettingactive larvaeinto beakers plored with fresh 1 um-filteredsea water.Subsamples of these munities (Gains and Roughgarden, 1985; larvae were preservedfor later use in 3% Formalin. Roughgardenet al., 1985). Currentstudies Larvaethat were inactive at the bottom of the bowls are examining factors that determine the were presumedunhealthy or dead. Such larvae were settlement rate such as currents and pre- usuallyalive but not feedingand were more prone to dation in the et bacterialinfection than the active larvae.To minimize plankton (Roughgarden al., the risk of bacterialcontamination, the unhealthylar- 1987; Roughgardenet al., 1988). Such work vae were discarded. requiresidentification of barnaclelarvae in Larvaeof Chthamalusare particularlyprone to be- mixed-species plankton samples. The pres- coming caught on debris due to their fine setulation ent was conducted to examine the and small size. Therefore,to rearthese larvaesuccess- study fully, all particulatecontaminates coming from the air morphologicaldifferences between the lar- or filterbags were removed regularlyand the alga was vae of C. fissus, not previously described, monitoredcarefully for evidence of clumping. and C. dalli, previously described from Analysis of Larvae.- Formalin-preservedlarvae were Russian materialby Korn and Osyannikova examinedunder a compoundmicroscope with a com- (1979). Electrophoreticand morphological bined phase contrast/Nomarskiattachment. Size was with shell determinedwith an ocular micrometer.Total length characterstogether distinguishing was measuredfrom the frontalmargin to the tip of the featuresare presented.We found that adult dorsalthoracic spine. Cephalicshield lengthwas mea- C. dalli and C. fissus can be separated by sured from its frontal marginto its posteriorborder, 242 MILLER ETAL.: COMPARISON OF LARVAE AND ADULTS OF CHTHAMALUS DALLI AND C. FISSUS 243 d A.: :j B ?i?2i; X r r- 4t *, '. A ;? :eT ...I :r- :t' is t?1 j? ? 100 jum I I Fig. 1. Photographs of the characteristic setae of the second cirrus of adult Chthamalus dalli and C. fissus. Setae of C. dalli (A) are finely bipectinate, while setae of C. fissus (B) are coarsely bipectinate and have a pair of basal guards (see arrow). and its width measured behind the frontolateral horns. dalli appear finely bipectinate (Fig. 1A), In addition, drawings were made with a camera lucida while the setae of C. are bi- attachment. At least 50 larvae of each species at each fissus coarsely stage were examined for the drawings. pectinate and have a pair of basal guards this setal characteristic to Adult Descriptions. -Adult C. dalli and C. fissus were (Fig. 1B). Using collected from the field to examine and to photograph separate the species initially, electrophore- the characteristic differences in the second cirrus, a sis showed that C. fissus and C. dalli can be feature noted by W. A. Newman (personal commu- clearly distinguished on the basis of their nication). In addition, shell features that might be used malate to separate the species in the field were quantified. dehydrogenase (MDH) enzymes (Fig. 2A). Chthamalus has two loci encoding Electrophoresis. -Adult barnacles, Chthamalus spp., malate and were collected, dissected, and identified to species by dehydrogenase (Mdh- Mdh-2, the second cirrus characteristic. Sixty barnacles of each see Figs. 2A, B); both enzymes are dimers, species (the sample of C. fissus included all three oper- so that the single-banded phenotypes rep- cular morphs) were then placed in individual wells in resent homozygotes and triple-banded phe- a and covered with a few of 0.5M plastic tray drops notypes represent Tris-HCL buffer, pH 7.1. Each barnacle was homog- heterozygotes. Although enized within the well with a glass pestle and the bar- both Mdh-1 and Mdh-2 are highly poly- nacle extract was absorbed with a filter-paper wick. morphic loci, the two species have different Horizontal starch-gel electrophoresis and enzyme- alleles (further work with larger samples has staining were carried out on the samples as described led to the identification of at least six alleles The by Hedgecock (1979). samples were then stained at each locus and un- for malate dehydrogenase enzyme. (Blower Hedgecock, published observations)). The difference in RESULTS the alleles is particularly clear for the Mdh- I Adult Descriptions locus (Fig. 2A), and served to confirm the Under microscopic observation (400 x), species classification using the setae on the the setae on the second cirri of Chthamalus second cirri. 244 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 2, 1989 A Mdh-2 (113) Mdh-2 - * (114) Mdh-1 (108) (100) Mdh-1 -- C. daill C. fissus B Mdh-2 *9::000000,00 0 }Mdh-2 { 0 ??0 S@0 000 ???*@ go??0* 0 0.... 0 0 0 Mdh-1 { 00 910,99 0 000000 }Mdh-1 i C.C dallida C. fissus Fig. 2. Results of gel electrophoresis on Chthamalus dalli and C. fissus. A. Horizontal starch gels stained for malate dehydrogenase (MDH) enzyme. The characteristic migration distances are different for C. fissus and C. dalli and hence this assay can be used for species identification. The single bands at both the Mdh-1 and Mdh-2 loci of both species indicates homozygosity. B. A drawing of a horizontal starch gel stained for malate dehy- drogenase (MDH) enzyme. Both species are extremely polymorphic at both loci (Mdh-l and Mdh-2). Both heterozygotes (3 bands) and homozygotes (1 band) at each locus are shown. External characters allow an approximate and a dark brown band at the base are C. distinction of C. dalli from C. fissus to be fissus (Fig. 3C-E). However, we found that made in the field. We have found that the 12% of adult Chthamalus spp. with oval two species can be identified in the field with orifices possessed a mixture of these char- approximately 90% accuracy, measured acteristics (19/156 of C. dalli and C. fissus against the setal characteristics, using a combined; Table 1). In C.
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