Alpheus angulatus, a new species of snapping shrimp from the Gulf of Mexico and northwestern Atlantic, with a redescription of A. heterochaelis Say, 1818 (Decapoda: Caridea, Alpheidae)

14 Pages·1995·6.2 MB·

by  M R Mcclure

PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
108(l):84-97. 1995.
Alpheus angulatus, a new species of snapping shrimp from the
Gulf of Mexico and northwestern Atlantic, with a
redescription of^. heterochaelis Say, 1818
(Decapoda: Caridea: Alpheidae)
Matthew R. McClure
Business, Science, & Math Division, Lamar University-Orange, 410 Front Street,
Orange, Texas 77630, U.S.A.
Abstract.—Two species ofthe Edwardsii-group ofsnapping shrimp, Alpheus
heterochaelis Say and A. estuariensis Christoffersen, have been recorded from
coastal estuarine habitats of the Gulf of Mexico and Atlantic coasts of the
United States. A new species, A. angulatus, has been discovered to inhabit
thesehabitatsacrossthisrangeandisdescribedandillustrated. Thenewspecies
is morphologically similar to A. armillatus, the latter being a tropical species.
A redescription with illustrations of^. heterochaelis is also provided herein.
The Edwardsii species group oiAlpheus Carolina and south Florida, and proposed
consists ofat least 10 western Atlantic spe- that^. heterochaeliscomprisedtwoormore
cies(Chace 1972; Christoffersen 1979, 1984; species. From a reevaluation ofAlpheusin-
Williams 1984; Abele & Kim 1986), of habitingshallow-estuarineenvironmentsof
which Alpheus heterochaelis Say, 1818, is the western Atlantic, Christoffersen (1984)
the most abundant and widely distributed. suggested the existence of a species com-
Alpheus heterochaelis was originally de- plex, and described a new species, A. estu-
scribed from Amelia Island, Nassau Coun- ariensis (holotype from the Rio Potengi es-
ty, Rorida, and has been reported to range tuary, Rio Grande do Norte, Brazil), rang-
from the lower Chesapeake Bay to Aransas ing from the east coast ofRorida into the
County, Texas, Cuba, Curasao, Surinam, GulfofMexico from Mississippi to Texas;
and possibly to Sao Paulo, Brazil (Chace Cuba; Dominican Republic; Trinidad; Cu-
1972, Williams 1984). rasao; and from Ceara to Parana, Brazil.
A variety ofstudies have been done con- Christoffersen(1984)furtherconcludedthat
cerning the biology oiA. heterochaelis (for the Alpheus occurring in the northern Gulf
example, Wilson 1903, Nolan & Salmon of Mexico represents A. estuariensis, and
1970, Knowlton 1973, Conover & Miller describedthe range of^. heterochaelistobe
1978,Mellon&Stephens 1978).Despitethe from North Carolina to Paraiba, Brazil.
abundance of studies on A. heterochaelis, However, usingChristoffersen's(1984)key,
thegeographiclimitsandthesystematicsta- specimens from Galveston were identified
tusofthisspeciesremainproblematic. Based bytheauthorasA. heterochaelis. Thus, plus
on apparent misidentifications ofWest In- a reexamination of museum specimens of
dianandBrazilianspecimens, Chace(1972) A. estuariensis from the northern Gulf of
questioned the occurrence of A. hetero- Mexico suggest the existence oftaxonomic
chaelis south ofSurinam. Knowlton (1973) ambiguity concerning the Edwardsii group
noted differences in egg size between labo- ofAlpheus from the coastal waters of the
ratory reared A. heterochaelis from North GulfofMexico and northwestern Atlantic.
VOLUME 108, NUMBER 1 85
An assessment ofallozymic variation of armillatus is not available; it is likely that
specimens ofAlpheus from Texas revealed the specimens have been lost. As the only
the existence oftwo discrete and markedly detailed descriptions ofA. heterochaelis are
different gene pools (McClure & Green- from the Carolinas (Christoffersen 1984),
baum 1994).Additionalallozymicdata(un- comparisons to this species were from ma-
published) indicate that these forms are terial from the type locality and from the
sympatric throughout the northern Gulfof Carolinas.For^. armillatus, specimensfrom
Mexico and Northwestern Atlantic coasts the type locality (Antilles) were used as a
as far north as Beaufort, North Carolina. reference. Type material ofthe two species
The present study was designed to provide described here have been deposited in the
morphological descriptions and identifica- USNM.
tions ofthe two electrophoretically identi-
fied species. Alpheus angulatus, new species
Figs. 1, 2
Materials and Methods Crangon armillatus.—Hay & Shore, 1918:
386, fig. 9. {notAlpheusarmillatusMilne-
Snapping shrimps were collected at low
Edwards, 1837).
tidebydipnetinintertidalandshallowsub-
tidalhabitatsconsistingofsandormudbot- Alpheus estuariensis.—ChristoffersQn, 1984:
191 (in part, see discussion).
toms covered with oyster clumps or rocks.
Alpheus armillatus.—Chsice, 1972:62 (in
Shrimps were collected in coastal waters
part, see discussion).
from south Texas to North Carolina (see
mm
Appendix for localities). Shrimps were Holotype.—MalQ, 28 TL, on mud
—
stored at 80°C to fully preserve color pat- underrocksandrubble, SouthPadreIsland,
terns. Starch-gel electrophoresis (McClure Texas, Laguna Madrejust north ofBrazos-
& Greenbaum 1994) was used to sort the Santiago Pass, coll. M. K. Wicksten, 4 Jul
USNM
individuals into discrete electromorphic 1992, 266804.
classes. Morphologicalcharacterswerethen Material examined.-See Appendix.
assessed and compared with museum ma- Diagnosis.—Rostro-orbital depressions
terial and descriptions in the literature. To- abrupt posteriorly. Ventral margin of car-
tal Length (TL) of specimens is the com- apacepronouncedatanangleventrallypos-
bined measurements of the carapace, ab- terior to second pereopods. Minor claw of
dominal, and telson lengths. malenotbalaeniceps-shaped. Spinepresent
Museum specimens of A. heterochaelis, onmerusoffirstpereopod.Thirdandfourth
A. estuariensis, A. nuttingi, and A. armil- pereopods with movable spine on ischium,
latus were obtained from the following in- lacking on fifth pereopod.
stitutions: Marine Research Division, Flor- Descriptionofholotype.—Rostrumreach-
ida Department of Natural Resources, St. ing 0.5 length of first antennular segment;
Petersburg, Florida (FBSC); Marine Envi- in form of raised crest extending beyond
ronmental Sciences Consortium, Tuscaloo- base ofeyestalks and widening into flat tri-
sa, Alabama (MESC); Texas A&I Univer- angulararea (Fig. 1A). Ocularhoods prom-
sity, Kingsville, Texas (TAI); National Mu- inentandunarmed, separatedfromrostrum
seum ofNatural History, Smithsonian In- by adrostral depressions abrupt posteriorly
stitution, Washington, D.C. (USNM); (Fig. lA). Ocular hood width 0.27 times
MuseumnationalD'Histoirenaturelle, Par- lengthofcarapace. Carapaceasfigured (Fig.
is, France (MNHN); LamarUniversity col- IC), 0.35 times TL (range 0.31-0.40 for all
lection, Beaumont, Texas (LU). The origi- specimens examined). Carapace smooth,
nal type material of^. heterochaelis and A. posterior of carapace with cardiac notch;
S6 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig. 1. Alpheus angulatus, new species. Adult male (holotype) from South Padre Island, Texas (TL = 28
mm). A, anterior carapace and antennae, dorsal; B, sixth abdominal somite, telson and uropods, dorsal; C,
carapaceandantennae,lateral; D, abdominalpleura, lateral. Barindicates 5 mm.
VOLUME NUMBER
108, 1 87
ventralmarginacuteventrallyjustposterior odusas figured (Fig. 2A, B), with upperand
to articulation ofsecond pereopod. lower notches forming saddle-like depres-
Abdomenasfigured(Fig. ID), 0.55 times sions (extending about 0.07 and 0.11 times
totallength (range 0.44-0.56). Pleura ofab- length of propodus, respectively) into the
dominal somites 1 through 5 with ventral lateral surfaces of propodus; upper notch
margins rounded. Sixth abdominal pleura positionedabout0.5thelengthofpropodus,
withventral margin acute posteroventrally. lower notch positioned about 0.61, having
Telson (Fig. IB) 0.11 times total length tuftofsetaeonproximalendoflowernotch;
(range 0.09-0.14). Proximal telson width angularareaofuppermesialsurfaceoffixed
0.67 of length, distal width about 0.5 of fingerrounded, havinggranulartexture and
length; 2 pairs of dorsal spines, anterior- tuftsofsetae; distalendofpropodusround-
most pair positioned about 0.4 of telson ed, with tufts ofsetae. Minor chela robust
length, posteriormost pair positioned at and setose, with setal tufts increasing dis-
0.67; posteriormargin convex, with 2 pairs tallyonbothpropodusanddactyl(Fig. 2C);
oflateral spines, space between spines with notsexuallydimorphic, lackingbalaeniceps
double row ofsetae. setose crest on dactyl; propodus 0.33 times
Antennular peduncle with stylocerite totalbodylength (range 0.15-0.37 forspec-
dorsally flattened, terminating anteriorly at imens examined); propodus height about
sharp point which reaches anterior margin 0.30timeslengthinbothsexes; dactyllength
of first antennular segment; second anten- about 0.5 times that ofpropodus length in
nular segment longer than first and about both sexes.
1.5 times as long as third (Fig. lA, C). Second pereopods slender and weakly
Antennal spine reaching end ofantennal chelate (Fig. 2D); carpus subdivided into 5
peduncle, and just overreaching antennal articles decreasing in length as follows
scale; concave atmiddle and faintly convex (numbered from the proximal end); 1, 2, 5,
at distal tip. Basal segment ofantennal pe- 3 = 4. Third and fourth pereopods with
dunclearmedwithspineventrolaterally(Fig. ventral movable spine on ischium (Fig. 2E,
lA, C). F); dactylisimple; thirdpereopodpropodus
Mandiblewith 1 teeth. Thirdmaxilliped with 7 stout spines and 3 or 4 smaller al-
reaching just beyond end of antennal pe- ternating lateral spines, fourth with 8 stout
duncle; terminal article setose. spines and 5 or 6 alternating lateral spines.
First pereopods (Fig. 2A, B, C) strongly Fifth pereopod (Fig. 2G) with 8 or 9 spines
chelateandunequal;merusarmedwithspine on the propodus and several lateral bands
distoventrally. Major chela thick, setose ofcomb-like setae extending on distal half;
distally; propodus length 0.51 times total ischium without spine. Second pleopod of
body length (range 0.28-0.56); upper and malewith appendix masculina shorterthan
lower margins deeply notched proximal to appendix interna (Fig. 2H).
articulation of dactyl, upper notch width Coloration.—OyevaW dark olive green to
about 0.04 ofpropodus length, lower notch brown, occasionallywithblue tonesaround
width about 0.1 1 ofpropodus length; max- orbits, distal half ofabdomen, and telson;
imum propodus height about 0.46 times bodyspeckledwithtinybrownorred-brown
length; dactyllengthabout0.36 timesprop- spots.Majorcheladarkolivegreenthrough-
odus length; hand height with dactyl closed out mesial surface, unspeckled; upper and
about 0.35 times propodus length; dactyl lowernotcheseithercoloredasrestormajor
withentireupperanddistal marginsround- chelaorlightenedtopaleyellow, rarelywith
ed, with setae arranged in tufts; opposable blue; inner surface ofmajor chela white or
margin ofdactyl with molar process tilted pale yellow, occasionally bluish; fingertips
at angle to axis ofdactyl; sculpture ofprop- paleyelloworwhite. Minorcheladarkolive
88 PROCEEDINGS OFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig. 2. Alpheus angulatus, new species. A, major first pereopod (right), dorsal; B, same, ventral; C, minor
firstpereopod(left);D,leftsecondpereopod;E,leftthirdpereopod;F,leftfourthpereopod;G,leftfifthpereopod;
H, left secondpleopod. Barindicates 5 mm.
,
VOLUME NUMBER
108, 89
green. Secondtofifthpereopodstranslucent Material examined.—See Appendix.
—
to reddish, lightly speckled with reddish Diagnosis. Rostro-orbital depressions
brown spots or mottled with red. Telson not abrupt posteriorly. Ventral margin of
colored as abdomen, highly speckled with carapace evenly rounded, not pronounced
tinybrown orred-brown spots; uropods of- atanangleventrallyposteriortosecondper-
ten bluish throughout, occasionally blue eopods. Minor claw of male balaeniceps-
distally. shaped. Spine absent on merus offirst pe-
Size.—Total length for males ranging 17 reopod. Third, fourth, and fifth pereopods
to 38 mm, females 17 to 35 mm. with movable spine on ischium.
Etymology.—Specificnamederivedfrom Description of neotype.—KosXruvcv cari-
the shape ofthe ventral margin ofthe car- nate, reaching 0.5 length offirst antennular
apace, which contains a pronounced angle segment, extending about as far as base of
posterior to the second pereopods. eyestalks. Ocularhoods prominent and un-
Range.—The known range for this spe- armed, separated from rostrum by shallow
ciesisthroughoutthenorthernGulfofMex- adrostral depression. Ocular hood width
ico and northwestern Atlantic as far north 0.25 times length ofcarapace (Fig. 3A, C).
asBeaufort,NorthCarolina, andasfarsouth Carapaceasfigured(Fig. 3C), 0.35 timesTL
as Quintana Roo, Mexico, and Haiti. (range 0.32-0.40 for all specimens exam-
Habitat.—IntQTtidal and shallow waters ined). Carapace smooth, posteriorwith car-
in bays and other quietwaters consisting of diac notch.
mud bottoms with oyster clumps or rocks Abdomenasfigured(Fig. 3D), 0.52 times
and rubble. totallength (range 0.45-0.55). Pleura ofab-
dominal somites 1 through 5 with ventral
Alpheus heterochaelis Say, 1818 margins rounded. Sixth abdominal pleura
Figs. 3, 4 with ventral margin acutely rounded pos-
teroventrally. Telson (Fig. 3B) 0.12 times
Alpheus heterochaelis Say, 1818:243.—
— totallength(range 0.09-0.14). Proximaltel-
Milne-Edwards, 1837:356. Kingsley
sonwidthabout 0.67 oflength, distalwidth
1878a:194, 1878b:329, 1879:417.-
Brooks & Herrick, 1892:376, pi. 2.- about0.5 oflength; 2 pairs ofdorsal spines,
anteriormost pair positioned about 0.5 of
Coutiere, 1910:485.-Verrill, 1922:76, pi.
telsonlength, posteriormostpairpositioned
22, figs. 1, 2, 4a-c; pi. 24 figs. 7, 7a; pi. almost 0.75; posterior margin convex, with
30, figs. 1-la, it, 2a-2e; pi. 33, figs.
2pairsoflateralspines,spacebetweenspines
1, 2.-Holthuis, 1959:102.-Williams, with double row ofsetae (Fig. 3B).
1965:66,fig. 54, 1984:95,fig. 65.-Chace, Antennular peduncle with stylocerite
1972:67.-Coelho&Ramos, 1972:148.-
dorsally flattened and terminating anteri-
Christoffersen, 1984:200, figs. 5-7.
Crangon heterochaelis. —Hay & Shore, orly at a sharp point which reaches anterior
margin offirst antennular segment; second
1918:386, fig. 8.-Schmitt 1935:144. antennular segment longer than first and
Holotype.—Not extant, Amelia Island, about 1.4 times as long as third (Fig. 3A,
Nassau County, Rorida. C).
— mm
Neotype. l male, 31 total length. Antennal spine reaching end ofantennal
Fort Saint George Inlet (8 miles south of peduncle, and just overreaching antennal
Amelia Island), Duval County, Florida, in- scale; spine faintly convex at distal tip,
tertidal habitat consisting ofhard mud and straight orslightlyconcave at middle. Basal
clumps ofoysters, coll. M. R. McClure and segment of antennal peduncle armed ven-
L. S. McClure, 25 May 1992, USNM trolaterally (Fig. 3A, C).
268646. Mandible with 9 teeth. Third maxilliped
90 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig. 3. AlpheusheterochaelisSay, 1818.A-L,Adultmale(neotype)fromFortSt.GeorgeInlet, 8milessouth
ofAmeliaIsland,Florida(TL= 31 mm). A,anteriorcarapaceandantennae,dorsal;B,sixthabdominalsomite,
telsonanduropods,dorsal;C,carapaceandantennae,lateral;D,abdominalpleura,lateral.Barindicates5 mm.
VOLUME 108, NUMBER 91
Fig.4. Alpheusheterochaelis.A,majorfirstpereopod(right),dorsal;B,same,ventral;C,minorfirstpereopod
(left); D, left second pereopod; E, left third pereopod; F, left fourth pereopod; G, left fifth pereopod; H, left
secondpleopod; I, Adult female from Beaufort, NC, minorfirst(left) pereopod. Barindicates 5 mm.
9
92 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
reachingendofantennalpeduncle;terminal Coloration. —Overall olive green to
article setose. brown, oftenwithblue tones around orbits,
First pereopods strongly chelate and un- distal half of abdomen, and telson; body
equal; merus unarmed distoventrally. Ma- speckled with tiny brown or red-brown
jorchela(Fig. 4A, B)thick; propoduslength spots.Majorchelaolivegreenwithpalearea
0.45 times total body length (range 0.29- atlowercenterofpalm;raisedareasofpalm
0.61); upper and lower margins deeply and opposable margin of fixed finger very
notched proximal to articulation ofdactyl, dark green; upper and lower notches pale
upper notch width about 0.07 ofpropodus yellow to pale blue; inner surface ofmajor
length, lower notch width about 0.10 of chelawhite to pale blue; fingertips pale yel-
propoduslength;maximumpropodusheight low or white. Minor chela olive green. Sec-
about0.45 timeslength; dactyllengthabout ond to fifth pereopods translucent to pale
0.35 times propodus length; hand height blue, lightly speckled with reddish brown
with dactyl closed about 0.32 times prop- spots. Telson colored as abdomen; exopod
oduslength; dactylwithupperdistalmargin ofuropodsblueondistal segment; endopod
rounded, opposablemarginwithmolarpro- ofuropods blue distally.
cess tilted at angle to axis ofdactyl; sculp- ^/z^.—Neotype total length 31 mm. To-
ture ofpropodus as figured, with upperand tal length for males ranging 16 to 45 mm,
lower notches forming saddle-like depres- females ranging 18 to 57 mm.
sions (extending0.07 and 0.14 times length Range.—l^o^Qx Chesapeake Bay south-
of propodus, respectively) into the lateral wardandwestwardintotheGulfofMexico;
surfaces of propodus; upper notch posi- Cuba; Curasao (Williams 1984); Bermuda
tioned about 0.5 the length of propodus, (Verrill 1922); Surinam (Chace 1972);
lower notch positioned about 0.67; distal southwardtoParaiba(Christoffersen 1984).
end ofpropodus rounded. Minorchela sex- Habitat.—InXQvXiddiX and shallow waters
ually dimorphic (Fig. 4C, I). Male minor in bays and other quietwaters consisting of
chela balaeniceps-shaped, having setose mud bottoms with oyster clumps or rocks
crest on dactyl and accessory' crest on op- and rubble.
posable margin ofthe propodus; propodus
0.32 times total body length (range 0.22-
Discussion
0.40 for specimens examined); propodus
heightabout0.24timeslengthinmales,0.1 Although Alpheus angulatus and A. het-
times length in females; dactyl length about erochaelisaremorphologicallysimilar, they
0.45 timesthatofpropoduslengthinmales, are not closely related. Alpheus heterochae-
and about 0.5 in females. lis and A. angulatus were referred to as Al-
Second pereopods (Fig. 4D) slender and pheusgroup A and group B, respectively in
weaklychelate; carpussubdividedinto 5 ar- McClure & Greenbaum (1994), where the
ticles decreasing in length as follows (num- twospeciesareshowntobehighlydivergent
=
bered from the proximal end): 1, 2, 5, 3 electrophoretically. Alpheus heterochaelis
4. Third to fifth pereopods (Fig. 4E, F, G) was the most common species that the au-
with ventral movable spine on ischium; thor collected from Texas and from North
dactyls simple; third pereopod propodus Carolina. Thisspeciesdiffersfromtheother
with9 or 1 stoutspines, fourthwith 8 stout Edwardsii-groupAlpheusmentionedherein
spines;fifthpereopodwith7spinesonprop- in that A. heterochaelis possesses a balaen-
odus, with several lateral bands of comb- iceps-typeminorclawinmales, anda mov-
like setae extending on distal half Second able spine on the ventral surface ofthe is-
pleopod of male with appendix masculina chium of the fifth pereopod. Alpheus het-
shorter than appendix interna (Fig. 4H). erochaelis is further distinguished from A.
VOLUME NUMBER
108, 93
angulatus in that the rostro-orbital area in the color pattern, the two species are diffi-
the former lacks posteriorly abrupt adros- cult to distinguish. Both A. armillatus and
tral depressions and lacks the broad trian- A. angulatushavethe typicalA. armillatus-
gular area ofthe carapace from which the type rostrum, consisting of a broad, flat-
rostrum terminates posteriorly. tened triangular area posteriorly, and pos-
Material of^. estuariensiswas examined teriorlyabruptadrostralfurrows.Milne-Ed-
from Port Isabel and Gilchrist, Texas, and wards' (1837) description of^. armillatus
from Mississippi Sound, Alabama, as well was brief and lacking in detail. However,
as the Gulf of Mexico material used by the locality was given as the Antilles. The
Christoffersen (1984). Alpheus estuariensis species was named "the banded Alpheus''
was not found in this survey, and thus is for its conspicuous banded color pattern;
probably of uncommon occurrence in the this pattern is lacking inA. angulatus. Ver-
northernGulfofMexico.Alpheusestuarien- rill (1922) stated that A. armillatus from
sis is distinguishable from A. heterochaelis Bermuda, when recently caught, was iden-
in that it lacks the balaeniceps-type minor tifiable by its conspicuous transverse bands
claw in males, and lacks the movable spine ofwhite on the body and rings ofcolor on
onthe ischium ofthe fifth pereopod. In ad- the legs and antennae, and provided a pho-
dition, the minor claw of^. estuariensis is tograph (plate 20, fig. 4b) showing such a
very slender relative to that ofA. hetero- pattern.Zeiller(1974, p. 76)providedacol-
chaelis. orphotographofA. armillatusrevealingthe
Alpheusangulatuswasthemostcommon conspicuous banded pattern on the abdo-
species that the author collected from Flor- men,withthebodyandchelaecoloredover-
idaandfromLouisiana, andhasapparently all brown speckled with white. In ^. an-
been mistaken forAlpheus heterochaelis in gulatus, thebodyisdarkerandspeckledwith
a number of museum collections. This is dark red-brown spots, with speckling lack-
becausetheyarebothsimilarlycolored, and ing on the chelae.
commonly taken in a single sampling (both Hay & Shore (1918) Hsted A. angulatus
havebeen collected simultaneouslyat most as A. armillatus from North Carolina. Al-
ofthe author's collecting localities). though the author was informed that Hay
In one case, Alpheus angulatus has been & Shore's (1918) collection may no longer
mistaken forA. estuariensis by Christoffer- exist(R. B. Manning, pers. comm.), thecol-
sen (1984). One of the specimens referred or description by Hay & Shore for the A.
to as A. estuariensis (USNM 98137) from armillatus from Beaufort, North Carolina,
Louisiana represents A. angulatus and not matches that of^. angulatus. In addition,
A, estuariensis. Alpheus angulatus differs the figures provided by Hay & Shore of^.
from A. estuariensis in that the former has armillatus matchA. angulatus with respect
amorerobustminorclaw,whereasthespine totheshapeoftherostrumandminorchela.
onthemerusofthe firstpereopodislacking They also mentioned the rarity ofthis spe-
in^. estuariensis. Inaddition,theupperand ciesatBeaufort,NorthCarolina, whichalso
lower notches ofthe major chela are wider confirms the authors' findings (one A. an-
of^. estuariensis than those of^. hetero- gulatus individual was caught with 54 A.
chaelis and of^. angulatus, and the distal heterochaelis individuals). The color pat-
end ofthe propodus is sharply truncated in tern difference between the Carolinian A.
A. estuariensis. armillatusandthetropicalA. armillatusalso
Alpheusangulatushaspreviouslybeenre- led Knowlton (1970) to speculate that the
ferredtoastheCarolinian^. armillatus(Hay Carolinian form may represent a separate
&Shore 1918;Knowlton 1970;Chace 1972; species.
Williams 1965, 1984). Exceptbyexamining The likelihood thatA. armillatus, as cur-