Table Of ContentPROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
108(3):458^64. 1995.
Within-species variation in Periclimenes yucatanicus
(Ives), with taxonomic remarks on P. pedersoni Chace
(Crustacea: Decapoda: Caridea: Palaemonidae)
Mary K. Wicksten
A&M
Department ofBiology, Texas University, College Station,
Texas 77843, U.S.A.
Abstract.—The spotted cleaner shrimp Periclimenesyucatanicus is found to
vary in the number of rostral teeth, length/width ratio of the carpus of the
major cheliped, color pattern and cnidarian host. Similar variation has been
foundin otherspecies oiPericlimenes. Morphological featuresandtype ofhost
&
have been used to distinguish between P. anthophilus Holthuis Eibl-Eibes-
feldt, andP. pedersoniChace, but these features overlap extensively; therefore,
P. anthophilus is herein considered to be ajunior synonym ofP. pedersoni.
Shrimp ofthe genus Periclimenes Costa, those of species ofPericlimenes known to
1844areamongthemostcolorfulcarideans inhabit sea anemones in the area. Records
ofcoral reefs. Species inhabiting cnidarian ofshrimpspeciesandtheirhostswerequan-
hosts are among the best known. Four spe- tified. The results indicate thatthere iscon-
cies ofPericlimenes have been reported to siderablewithin-speciesvariationinatleast
livewithcnidariansinthewesternAtlantic, two species oiPericlimenes.
GulfofMexico and Caribbean: P. yucatan-
icus (Ives 1891), P. pedersoni Chace, 1958
Methods
&
P. anthophilus Holthuis Eibl-Eibesfeldt,
1964 and P. rathbunae Schmitt, 1924. (See During studies ofcleaning behavior, this
Chace 1972, fora key to the species oiPer- writer and volunteer divers from the non-
CEDAM
iclimenes in the area). profit organization International,
During studies of cleaning behavior of counted numbers ofshrimp per species of
shrimpandfish,specieso{Periclimeneswere host in Key Largo, Florida, U.S.A., during
observed on coral reefs near Key Largo, 1 1-17 July 1992, andBonaire, Netherlands
Florida, U.S.A., and Bonaire, Netherlands Antilles, during25 July-5 August 1993 and
Antilles. The shrimp were identified from 30 July-12 August 1994. Shrimp were ob-
photographs and by sight. However, during servedduring SCUBAdives atdepthsof3-
m
night dives at Bonaire, shrimp with an un- 22 between 0600-2200 hours. Twenty-
usual color pattern, unlike that of known four photographs of all species of Pericli-
species in the area, were observed. These menes were taken in Florida and 1 1 in
shrimp were found in association with the Bonaire. Species were identified in the field
giant sea anemone, Condylactis gigantea accordingtothecolorpatterns showninthe
(Weinland). popular book by Humann (1992:151); un-
To identify this unknown shrimp, speci- identified ones were reported as Pericli-
mens were collected and compared with menes sp.
previouslyidentifiedmaterial. Photographs Five specimens (the maximum number
oflive animals were examined to compare allowed as stated on the collecting permit)
color patterns ofthe unknown shrimp with ofthe unidentified shrimp were collected at
VOLUME NUMBER
108, 3 459
Fig. 1. Variation imnmtherostrum and second pereopodofPericlimenesyucatanicus. a, left seconmdmpereopod
fromindividual29.8 intotallength;b,c,leftandrightsecondpereopodsfromindividual 16.5 intotal
mm mm
length;d,rostrumfromindividual29.8 intotallength;e,rostrumfromindividual 13.0 intotallength.
mm
Scalesare 1 and referto thedrawings immediatelyabovethem.
Calabas Reef, near Kralendijk, Bonaire, previouslycollected material ofthe species,
during 1994. An additional 13 were pho- there is variation in the number ofrostral
tographed in their natural habitat, but not teeth and the length-width ratio ofthe car-
collected. The animals were kept alive for pus ofthe second (major) cheliped (Fig. 1).
5-7 days and their color patterns recorded. The rostrum can have 5-8 dorsal and 0-3
These specimens were compared with de- ventral teeth in addition to an acute apex.
scriptions of known species and also with The length/width ratio ofthe carpus ranges
13 specimens of P. yucatanicus from the from 2.3-7.5. A more slender carpus (4x
collectionsoftheNational Museum ofNat- longerthan wide ormore) usually occurs in
mm
ural History Smithsonian Institution, smallershrimp(total length 16 orless).
mm
Washington, D.C. The color patterns also (The largest specimen measured 29.8
were compared with those shown in pho- in total length, orbit-telson.) In three spec-
tographsofP.yucatanicusfromGrandCay- imens, the second pereopods were asy-
man, British West Indies (two slides show- metrical.
ing three shrimp), Isla Cozumel, Quintana The 18 shrimp of the nocturnal color
Roo, Mexico (three slides showing two phase had translucent bodies with a few
shrimp), the U.S. Virgin Islands (one pho- small spots ofbrown orwhite on the dorsal
tograph ofone shrimp) and Discovery Bay, surface, and a translucent turquoise blotch
Jamaica (photograph by Colin 1978). near the gastric region. In two, there was a
pale white stripe along the dorsal midline;
inanothertwo,therewasasmalldorsalpale
Results
brown spot on either the carapace or third
The unidentified shrimp are a previously abdominal somite. The pereopods were
unreportedcolorvariety(hereaftercalledthe banded. Most had white antennae, but two
nocturnal color phase) of P. yucatanicus. had the outer antennal flagella banded and
Their bodies closely match the description the inner ones white. In two, the uropods
ofP.yucatanicusasgivenbyHolthuis(1951: ended in faint white spots; in the others,
38, pi. 10, figs. a-1). In these specimens and there either were no such spots or the tail
1
460 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig.2. ColorvariationinPericlimenesyucatanicus,drawnfromphotographsinHfe.a,entireanimal,showing
banded appendages, from Florida, day; b, individual from Bonaire, day; c, individual from Bonaire, night; d,
individual from Bonaire, day; e, same individualas seenindrawingd, night; f, individual from Bonaire, day.
fan could not be observed in the photo- the 82 photographs identifiable as P. yu-
graph. catanicus taken at Bonaire, all showed
Twooftheindividualsthatwerecollected shrimp with the pereopods banded with
at night are shown in the nocturnal color whiteandacontrastingdarkershade ofred,
phase in Figs. 2c and 2e. By day, both purple or violet. Thirty-two shrimp (39%)
changed their color pattern to that oflarge had entirely white antennae; 34 shrimp
white, tan, green or purple blotches, spots (41%) had the outer flagellabanded and the
and saddle marks on a translucent body. innerwhite, and 16 shrimp (20%) had both
The same individual is shown in Figs. 2d antennal flagella banded. Eleven photo-
and 2e, illustrating the marked change in graphs showed onlythe front ofthe animal,
colorpattern. Examination ofthe live spec- and thus only the color ofthe appendages
imens and photographs revealed consider- andantennae could be seen. In the other 7
ablecolorvariationinP. yucatanicus. Some photographs, at least the carapace and first
of the patterns are shown in Figure 2. Of threeabdominal segmentswerevisible. The
VOLUME 108, NUMBER 3 461
Fig. 3. Periclimenespedersoni2i\ Bonaire.
color pattern typically included a large dor- of the shrimp were translucent except for
sal ''saddle" of pure white or pink, tan or the bands and spots. Similar color patterns
green with a white borderon thedorsal sur- were observed in photographs of shrimp
face ofthe carapace, (seen in 31 shrimp, or from otherareas except forone photograph
44% ofthe total), a saddle (sometimes in a from Grand Cayman, which showed two
figure-eight pattern) ofsimilar color on the shrimp with the same color pattern as the
dorsal surface ofthe first and/or second ab- nocturnal color phase from Bonaire. The
dominal somite (seen in 30 shrimp, or 42% photograph of P. yucatanicus by Humann
ofthe total), and a large saddle on the third (1992) shows a shrimp with a brown back-
abdominal somite (seen in 46 shrimp, or ground pigmentation.
65% ofthe total). The tips of the uropods Another common shrimp, Pcriclimenes
oftenweremarkedwithwhiteanddarkspots pedersoni, hadverydifferentmarkingswhich
shaped like eyespots; however, the tail fan were not observed to change at night. This
was visible in only 21 photographs. The specieswas marked with white, blueorpur-
lower surface of the carapace or abdomen ple-violet lines running the length of the
often had spots ofwhite or white and dark body, similarmarkson the majorchelipeds,
pigment and the anterior parts ofthe ceph- purple speckles on the maxillipeds and oth-
alothoraxcould bespeckled; however, it was erpereopods, and pure whiteantennae. The
difficult to see these marks in many pho- white lines and antennae are readily visible
tographs. Many shrimp also had "saddle" at a distance (Fig. 3).
marks on the fourth abdominal somite, or Of 302 total P. yucatanicus observed in
had marks that overlapped two abdominal thewild(whetherphotographedornot), 283
somites. Two of the largest shrimp (total were among the tentacles o{Condylactisgi-
body length approximately 30 mm, as es- ganlca, 13 were on the sea anemone Bar-
timated from photographs) had a body iholomea annulata (Lesueur) and six were
backgroundcolorofbrownishpigment; most away from a cnidarian host. Usually, only
462 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
one shrimp was observed per host, but as 1991:fig. 1;Colin 1978:344as"unidentified
many as five could be found per anemone. species oi^Periclimenes^^). It is possible that
All ofthe unidentified shrimp at night were slight regional differences in color pattern
found on C. gigantea. Periclimenes peder- may exist in these species, but such differ-
soni was more abundant. Ofa total of938 enceswouldbedifficulttoseeorphotograph
C
observed, 555 were on gigantea, 315 on in the natural habitat.
B. annulata and 68 on other hosts or away Marked changes in color patterns from
from a cnidarian. As many as 13 P. ped- day to night have been reported previously
ersoni could be found living in a single sea in the hippolytid shrimp Hippolyte varians
anemone. The two species never coexisted Heptacarpus pictus and Heptacarpus palu-
on the same host. The hippolytid shrimp dicola (Green 1961, Bauer 1981). In these
Thoramboinensis(De Man) often occupied species, the shrimp showed a tranluscent
C
the sameindividual of giganteawith one blue color by night, regardless ofthe color
ofthetwo species; snapping shrimp {Alphe- patterndisplayedbyday.DiversatKeyLar-
us?armatusRathbun)livedalongthestalks go noticed a similar color change in Thor
of^. annulataunderthetentacleswherethe amboinensis, which had a pattern ofwhite
Periclimenes lived. spotsagainstachocolatebrownbackground
No individuals ofP. rathbunae were ob- by day, and a blue background at night.
served at either Key Largo or Bonaire. Thereare fewobservationsofthenocturnal
colorationofshrimp,especiallyintheirnat-
ural habitat, so it is difficult to determine
Discussion how common diurnal/nocturnal color
Holthuis (1951) noted variation in the changes are among species oiPericlimenes.
length/width ratio ofthe carpus ofthe sec- Spotteetal. (1991)reviewedassociations
ond pereopod in specimens ofP. yucatan- ofshrimpwithcnidariansintheWestIndies
icus, and speculated on whether this might and Bermuda, and reported that neither P.
be due to sexual difference, a growth form yucatanicus nor P. pedersoni was confined
or geographic variation. This asymmetry to a single host. Periclimenes yucatanicus
also could be due to loss and regeneration was listed to live with six species of sea
ofanappendage. Theresultsgivenheresug- anemones (Order Actiniaria), the jellyfish
gest that variation in the proportions ofthe Cassiopeia xamachana (Bigelow) and the
carpusofthe second pereopodexistswithin corallimorpharian, Rhodactis sanctaetho-
P. yucatanicus. Holthuisnotedsimilarvari- masae(Duchassaing & Michelotti). Pericli-
ation in P. infraspinis (Rathbun). Chace & menes pedersoni was reported to live with
Bruce(1993)notedthatseveralIndo-Pacific six species of sea anemones, the same jel-
species of Periclimenes had subequal to lyfish and a tube anemone, Cerianthus sp..
grossly unequal second pereopods. At Key Largo, diver Rick Sammon photo-
The pattern of spots and lines in speci- graphed Periclimenes yucatanicus in asso-
mens from Bonaire and Florida was con- ciationwithanothercorallimorpharian,Ri-
sistent with that shown in photographs of cordeaflorida (Duchassaing & Michelotti).
P. yucatanicus from other areas. Less vari- I photographed the same association at
ation was seen in the color pattern of P. Grand Cayman.
pedersoni (Humann 1992:151; Sefton & Evidently, P. yucatanicus and P. peder-
Webster 1986:77, figs. 1 17, 1 18; Colin 1978: soni can live on multiple hosts. However,
339-341; Voss 1980:87 and color plate). most records of either species come from
Periclimenes rathbunae has a pattern of the anemones C. gigantea and B. annulata.
white and brown dots and lines, with white Goy (1990) found that more larval P. yu-
dots at the tips ofthe uropods (Spotte et al. catanicus completed metamorphosis sue-
VOLUME NUMBER
108, 3 463
cessfuUy if exposed to exudates of C. gi- well aswith otherhosts(Spotte etal. 1991).
ganteathantothoseofotherspeciesornone Lacking additional evidence of distinctive
atall; larvalP.pedersonishowedbettersur- coloration or species-specific behavioral or
vival ifexposed to exudates of^. annulata. morphological distinguishing features, P.
&
Levine Blanchard (1980) reported that anthophilus should be considered a junior
species of Periclimenes could be stung by synonym ofP. pedersoni Chace, 1958.
sea anemones to which they were not ac-
climated. Acknowledgments
&
EibPeesrfiecllditm(e1n9e6s4a)ntwhaosphdielsucsriHboeldthausiasdisEtiibnlc-t MaIrianmegPraartekf,ulantodKtahleligdoeveMrenyemre,ntBoonfaitrhee
species from Bermuda. The original au-
NetherlandsAntilles forpermissionto con-
thors, and Chace (1972) remarked that it duct studies and collect specimens. Vol-
was very similar to P. pedersoni, but could CEDAM
unteer divers from International,
be distinguished by the position ofthe he-
especially Rick Sammon, Dave Downs and
patic spine in front of the most posterior
Stella Covre, and Nancy Sefton ofthe Cay-
dorsalcarapace spine; thecarpus ofthe sec- man Islands aided immensely in photo-
ondpereopodbeinglessthanhalfthelength
graphing and documenting shrimp in the
ofthe chela and the host beingActinia ber-
natural habitat.
mudensisor C. gigantea. Othermorpholog-
ical features and the color pattern were in-
distinguishable from those ofP. pedersoni. Literature Cited
Chace(1972)remarkedontheclosemor- Bauer, R. T. 1981. Color patterns ofthe shrimps
phological similarity between the two sup- HeptacarpuspictusandH.paludicola(Caridea:
posed species, and re-examined specimens. Hippolytidae).—MarineBiology 64:141-152.
Theonlydifferenceshefoundbetweenthem Chace, F. A., Jr. 1958. A new shrimp ofthe genus
Periclimenes from the West Indies.—Proceed-
were the "different habits" and the "pro-
ingsoftheBiologicalSocietyofWashington71:
portionatelyshortercarpusofthemajorsec- 125-130.
ond pereopod". However, proportions of . 1972. The shrimpsofthe Smithsonian-Bre-
the carpus ofthe second pereopod vary in din Caribbean Expeditions with a summary of
species ofPericlimenes as well as in other the West Indian shallow-water species (Crus-
tacea: Decapoda: Natantia).—Smithsonian
palaemonids. In species of freshwater
ContributionstoZoology 98:1-179.
shrimpofthegenusMacrobrachium, forex- , & A. J. Bruce. 1993. The caridean shrimps
ample, the entire second pereopod is mark- (Crustacea: Decapoda) oftheAlbatross Philip-
edly longer and more robust in adult males pineexpedition 1907-1910,part6:superfamily
than in juveniles or females. As shown in Palaemonoidea.—Smithsonian Contributions
toZoology 543:1-152.
Fig. 1, the length/width ratio ofthe carpus Colin, P. L. 1978. Marine invertebrates and plants
can vary in specimens ofPericlimenesspp., ofthelivingreefT.F.H. Publications,Neptune
as does the length ofthe carpus relative to City, NewJersey, 512 pp.
the chela. Costa,O.G. 1844. SuduenuoviGeneridiCrostacei
Although species ofPericlimenes appar- Decapodi Macrouri Nota.—Annalidelle Acca-
demiadegliAspirantiNaturalisti,Napoli2:285.
ently have preferred hosts, they are not re- Goy,J. W. 1990. Componentsofreproductiveeffort
strictedto a single species ofcnidarian. The and delay oflarval metamorphosis in tropical
review by Spotte et al. (1991) shows this marine shrimp (Crustacea: Decapoda: Caridea
lackofspecificity, asdoesa studyofshrimp and Stenopodidea). Unpublished Ph.D. disser-
A&M
living with sea anemones in central Japan tation,Texas University,CollegeStation,
(Suzuki&Hayashi 1977). BothP.pedersoni Green,TJe.xas1,9611.77App.biology ofCrustacea. H.F. & G.
and P. antChophilus have been reported to Witherby, Ltd., London, 180 pp.
live with gigantea and B. annulata, as Holthuis, L. B. 1951. A general revision ofthe Pa-
464 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
laemonidae (Crustacea Decapoda Natantia) of KennisderFaunavan Curacao. Resultaten Ee-
theAmericas.I.ThesubfamiliesEuryrhynchin- ner Reis van Dr. J. van der Horst in 1920.—
inae and Pontoniinae.—Allan Hancock Foun- Bijdragen Tot de Dierkunde Genootschap Na-
dation Occasional Papers 11:1-332. turaArtis MagistrateAmsterdam 23:61-81.
, & I. Eibl-Eibesfeldt. 1964. A new speciesof Sefton, N., &S. Webster. 1986. Afieldguideto Ca-
thegenusPericlimenesfrom Bermuda (Crusta- ribbean reef invertebrates. Sea Challengers,
cea, Decapoda, Palaemonidae).—Senckenber- Monterey, California, 112 pp.
giana Biologica 45:185-192. Spotte,S.,R.W.Heard,P.M.Bubucis,R.R.Manstan,
Humann, P. 1992. Reef creature identification. & J. A. McLelland. 1991. Pattern and color-
Vaughan Press, Orlando, Rorida, 320 pp. ationoiPericlimenesrathbunaefromtheTurks
Ives, J. E. 1891. Crustacea from the northern coast andCaicosIslands,withcommentson hostas-
ofYucatan, the harbor ofVera Cruz, the west sociationsinotheranemoneshrimpsoftheWest
coastofFloridaandtheBermudaIslands.—Pro- Indies and Bermuda.—GulfResearch Reports
ceedingsoftheAcademyofNaturalSciencesof 8:301-311.
Philadelphia(1891):176-207. Suzuki,K.&K.Hayashi. 1977. Fivecarideanshrimps
Levine, D. M., & O. J. Blanchard, Jr. 1980. Accli- associated with sea anemones in central Ja-
mationoftwospeciesofthegenusPericlimenes pan.—Publications ofthe Seto Marine Biolog-
to sea anemones.—Bulletin ofMarine Science ical Laboratory 24:193-208.
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