Table Of ContentG. S. Herbert & R. W. Portell, 2002 Page 307
to the early fossil taxa, are generic only. Attiliosa aenig-
ma differs from the undescribed early Oligocene species
from France in having a less angulate and sloped body
whorl, a broader parietal shield, heavier spiral ornamen-
tation, and a strongerposteriorchannel.Attiliosaaenigma
differs from the next earliest New World species,Attiliosa
gretae Yokes, 1999, ofthe late early Miocene ofFlorida,
in having a less angulate body whorl, a broader parietal
shield, heavier spiral ornamentation, spiral cords ofequal
rather than unequal strength, a weaker anal channel, and
a less recurved siphonal canal.
The new species superficially resembles Panamurex
rutschi Yokes, 1992 (Yokes, 1992: pi. 11, figs. 1-4) from
the Pliocene Punta Gavilan Formation of Yenezuela, par-
ticularly in the morphology of the axial ribs, apertural
lirae, and columellar nodules; however, P. rutschi differs
in lacking the prominent anal channel and in having
thicker, unpaired, and more widely spaced cords on the
body whorl. Older species ofPanamurex, particularly the
Paleogene and early Neogene species, all have open
spines on the body whorl and siphonal canal, and thus
are very different from A. aenigma, which lacks spines
altogether.
The paired condition of the spiral cords in A. aenigtna
and A. bozzettii is noteworthy because the occurrence of
similar spiral ornamentation in anumberofspecies ofTak-
4Fi5g4ur(eHo3l.otyAptet)i;lihoesiaghbtozz1e7t.t0iimmHo,uamrat,xi1m9u93m.dIiaRmSeNtBerI1G02.71.8m7m3./ ciiaes(,seDeeanbnovoem)u,rienxclu(dTiankgiao)nceooofkeiitsMgeaocloNgeiiclalMlSy olindesYtoksepse,-
Locality: Ras Hafun, Somalia, 150-200 m. a. Apertural view. b. 1975, may indicate a closer phylogenetic relationship be-
Abapertural view. (Photographs courtesy ofR. Houart) tweenAttiliosa and Takia than previously recognized. Un-
til now. Attiliosa has been compared only to Panamurex
or Calotrophon (Yokes, 1971, 1976, 1992, 1999). Detailed
2, 5, 7) and at least in one species ofCalotrophon (Yokes, studies of the ontogeny of this character and cladistic
1992: pi. 19, fig. 11), but no clearly delineated posterior methods are necessary to determine whether the paired
channels are formed in species of Panamurex or Takia. condition is homologous in these different groups. We
Despite this consistency, we regard our generic place- draw attention to this condition primarily because it has
ment as tentative due to the low number of potentially been ignored in past species descriptions and systematic
informative characters in the type material. This problem reviews in the literature.
is attributable, in part, to poorpreservation, since the pro- A second fossil muricid (Figure 4) collected from Ter-
toconch and early teleoconch whorls are missing in the ramar 01 could be referable to A. aenigma because ofits
holotype. More problematic, however, is the relatively nearly identical shell shape, size, and paired spiral cords.
simple morphology of the new species, a condition that However, the axial ribs of this second specimen are nar-
characterizes a number of sub-lineages within the Poiri- rower, and the columellar nodules are more prominent
eria clan and has been a prime source of systematic con- than in the holotype of A. aenigma. Additional material
fusion in the Muricinae (Yokes, 1992, 1999). Strength- is needed to determine whether this specimen should be
ening our position somewhat is the close morphological included in A. aenigma or whether it represents yet an-
resemblance ofA. aenigma to the \W\ngAttiliosabozzettii other undescribed species.
Houart, 1993 (Figure 3) from deep waters off the coast The discovery ofA. aenigma has significance for our
of Somalia. Both A. aenigma and A. bozzettii exhibit understanding of the biogeographic history of the genus.
paired (or bisected) cords, a rounded rather than a shoul- Although Yokes (1989, 1992. 1999) proposed that the
dered body whorl, and up to four rather than only three genus originated in the Old World during the Paleogene
columellar nodules, although these characters vary some- and migrated westward in post-Paleogene times, the latest
what inA. bozzettii (Roland Houart, 2001, personal com- Eocene/earliest Oligocene age ofA. aenigma and its oc-
munication). These species differ in the more pronounced currence in the New World questions this interpretation.
posterior channel in A. aenigma. Although age resolution of the European material and
Similarities to other species ofAttiliosa. however, even sampling resolution ofthe Paleogene fossil record are too
———
Page 308 The Veliger, Vol. 45, No. 4
Berggren, W. A., D. V. Kent, C. C. Swisher, m, M.-R Aubry
& D. V. Kent. 1995. ArevisedCenozoicgeochronologyand
chronostratigraphy. Pp. 129-212 in W. A. Berggren (ed.),
Geochronology, Time Scales and Global Stratigraphic Cor-
relation. SEPM Special Publication 54.
Brewster-Wingard, G. L., T. M. Scott, L. E. Edwards, S. D.
Weedman & K. R. Simmons. 1997. Reinterpretation ofthe
peninsularFloridaOligocene: Anintegratedstratigraphicap-
proach. Sedimentary Geology 108:207-228.
Bryan, J. R. 1991. Stratigraphic and paleontologic studies of
Paleocene and Oligocene carbonate facies of the eastern
coastal plain. Doctoral Dissertation, The University ofTen-
nessee, ix -I- 324 pp.
Cooke, C. W. 1945. Geology ofFlorida. FloridaGeological Sur-
vey Bulletin 29:1-339.
Cooke, C. W. & W. C. Mansfield. 1936. Suwannee Limestone
of Florida (abstract). Geological Society of America Pro-
ceedings for 1935. Pp. 71-72.
Huddlestun, p. F. 1993. Arevision ofthe lithostratigraphic units
of the Coastal Plain of Georgia. The Oligocene. Georgia
Geologic Survey Bulletin 105. ix -I- 152 pp.
Jones, D. S., P. A. Mueller, D. A. Hodell & L. A. Stanley.
1993. *^Sr/'*''Sr geochronology of Oligocene and Miocene
marine strata in Florida. Pp. 15-26 in V. A. Zullo, W. B.
Harris, T. M. Scott & R. W. Portell (eds.). The Neogene of
Florida and Adjacent Regions. Proceedings of the Third
Bald Head Island Conference on Coastal Plains Geology.
Florida Geological Survey Special Publication 37.
Mansfield, W. C. 1937. Mollusks of the Tampa and Suwannee
limestones of Florida. Florida Geological Survey Bulletin
Figure 4. Attiliosa sp. cf. A. aenigma Herbert & Portell. UF 15:7-334.
104450; height 19.0 mm, maximum diameter 11.3 mm. Locality: Mansfield. W. C. 1939. Note on unreported Oligocene in Citrus
Terramar 01 (PO017), Suwannee Limestone, Polk County, Flor- County, Florida. Journal oftheWashington AcademyofSci-
ida, a. Apertural view, b. Abapertural view. ences 29(2):45-46.
Petuch, E. J. 1997. A new gastropod fauna from an Oligocene
back-reeflagoonal environment in west central Florida. The
poor to determine when and whereAttiliosa first evolved, Nautilus 110(4):122-138.
the timing and geographic position of the new species Randazzo, a. F. 1972. Petrography ofthe SuwanneeLimestone.
indicates, at the very least, that diversification and geo- Florida Geological Survey Bulletin 54 (part 2). 13 pp.
graphic range expansion inAttiliosa were occuning much VOKES, E. H. 1971. The geologic history of the Muricinae and
the Ocenebrinae. The Echo 4:37-54.
earlier than previously thought. Future studies should
VoKES, E. H. 1975. Cenozoic Muricidae ofthe western Atlantic
concentrate on refining the systematics ofPaleogene Mur-
region, pt. 6 Aspella and Dermomurex. Tulane Studies in
icidae from the Old World. Are there additional unde- Geology and Paleontology 11(3):121-162.
scribed or "lost" taxa referable toAttiliosa and/orclosely VoKES, E. H. 1976. Cenozoic Muricidae ofthe western Atlantic
related groups, and, ifso, what do they tell us about char- region, pt. 7 Calotrophon andAttiliosa. Tulane Studies in
acter evolution and biogeographic patterns within the Geology and Paleontology 12(3):101-132.
Muricinae? VoKES, E. H. 1988. Muricidae (Mollusca: Gastropoda) of the
Esmeraldas beds, northwestern Ecuador. Tulane Studies in
Acknowledgments. Rick Carter, Frank Garcia, Gary Morgan, Geology and Paleontology 21(1):1-50.
Craig Oyen, Kevin Schindler, and Thomas Stemann assisted VoKES, E. H. 1989. Neogene paleontology in the northern Do-
Rf.rWo.mP.Teirnracmoallrec0t1i.ngParotriaalcqfuuinrdiinnggsopfecfiiemlednwsorakndw/aosr pbruolvkidmeadtrbiyx minican Republic. 8. The FamilyMuricidae (Mollusca:Gas-
the Florida Museum of Natural History's McGinty Endowment. tropoda). BulletinsofAmericanPaleontology97(332):5-94.
VoKES, E. H. 1992. Cenozoic Muricidae ofthe western Atlantic
Geerat Vermeij and Roland Houart provided helpful discussions
oFlformiudraicCiodntsryisbtuetmiaotnictsoaPnadleboiboigoelooggryapnhuym.beTrhi5s18i.s University of rCeagliootnr,oppth.on9. AcPatnertyhnooltaubsi,a,PoainrdieArtitai,liAosspae:llaad,ditDieornmsoamnudrecoxr,-
rections. Tulane Studies in Geology and Paleontology 25(1—
LITERATURE CITED 3):1-108.
VOKES,E. H. 1999. Anotherlook atthe muricine gtnasAttiliosa.
Berggren, W. a., D. V. Kent, J. J. Flynn & J. A. Van Cou- The Veliger42(4):289-305.
VERING. 1985. Cenozoic geochronology. Geological Society VoKES, E. H. & A. D'Attilio. 1982. Review of the muricid
of America Bulletin 96:1407-1418. genus Attiliosa. The Veliger 25(1):67-71.
THE VELIGER
© CMS. Inc., 2002
The Veliger45(4):309-315 (October 1, 2002)
Latitudinal Gradients in Body Size and Maturation of
Berryteuthis anonychus (Cephalopoda: Gonatidae) in the
Northeast Pacific
JOHN R. BOWER', JAMES M. MURPHY- and YASUKO SATO'
'Faculty of Fisheries, Hokkaido University, 3-1-1 Minato-cho, Hakodate, Hokkaido 041-8611, Japan
-Auke Bay Laboratory. Alaska Fisheries Science Center, National Marine Fisheries Service, NCAA,
11305 Glacier Highway. Juneau. Alaska 99801-8626. USA
Abstract. Trends in body size and maturation with latitude of the gonatid squid Berryteuthis anonychus in the
northeast Pacific are described. Squid were collected during May 1999 at seven stations along 145° and 165°W between
mm
39° and 49°N. Mantle lengths ranged from 10.3 to 102.2 and increased significantly in both sexes from south to
north. Females were both larger and more numerous than males at the northern stations. Both sexes showed a clear
pattern of increasing maturity from south to north, and at each station, males were generally in a more advanced stage
of maturity than females. Most mature males occurred at the northernmost stations. No mature females were collected.
Our data suggest that B. anonychus migrates northward in the northeast Pacific during spring, with males maturing at a
smaller size than females.
INTRODUCTION stations along 145° and 165°W between 39°01' and
49°03'N (Figure 1). Sampling was conducted fromjust be-
Most studies of squid migration have been conducted on fore dawn to just after dusk. Both longitudinal transects
cshoomrmee.rScioamlelyofimtphoesretasnptecsipeesc,iessucthhaatsoTcocduarrroedleastipvaeclyifniecaurs nwoerrteh;s1a6m5p°leWd—innortdihffteoresnotuthd)i,reacntidontsher(e14w5as°Wn—osdoieulthpatt-o
pacificus (Steenstrup, 1880), Dosidicus gigas (d'Orbigny.
tern to the times at which stations were occupied, thus
1835), and Ille.x illecebrosus (Lesueur, 1821), have been eliminating the possibility that any trends in body size and
shown to migrate over long distances (> 1500 km) be-
maturation with latitude seen in the data might have been
tween low-latitude spawning grounds and high-latitude
feeding grounds (Hanlon & Messenger. 1996). Few stud- due to sampling bias. At each station, a midwater trawl
imeos,rehoowceevaenric, shpaevceiese.xamined the migration patterns of tmroadwilfiweadsto19f8ishmatlotnhge wsiutrhfacheexwaagsontaolwmeedsfhorin1whionugr.sTahned
smaBlelrr(ymtaeuntthliesleanngotnhyctohu1s50Pemamr)c,y o&ceaVnoiscs,squ1i9d63d,istirsiba- Tbordayw,liangndspaee1d.s2 cwmereme7s—h9lkinmerhwra',smaunsdedthien athveercaogdeennde.t
dimensions while fishing were 16 vertical spread and
uted mainly in the northeast Pacific (Roper et al., 1984). m
It is a major prey for salmonids (Pearcy et al.. 1988). 45 horizontal spread. Subsamples of the total catches
Pacific pomfret {Bramajaponica Hilgendorf, 1878; Pear- were taken at the two stations (#11 and #31) where more
cy et al., 1993), and neon flying squid {Ommastrephes than 400 squids were collected. A total of 359 B. anony-
chus specimens, including 195 males and 164 females,
bartramii (Lesueur. 1821); Pearcy, 1991). Despite its im-
portance in the food web of the subarctic North Pacific, were examined in the following analyses. Specimens were
liintftleereinsckesnoawrenmaabdouetointstlhiefemhiigsrtaotriyo.nInantdhesppraewsneinntgstoufdBy., Asexmedo,diafnidedthveedrosrisoanlomfantthleemlaetnugrtihtoyfsecaaclheswadsesmceriabseudrebdy.
anonychus in the northeast Pacific based on trends in Lipinski & Underbill (1995) was usedformaturity analysis
body size and maturation with latitude. (Table 1). The buccal membrane and inner mantle wall of
each female were examined for the presence ofdischarged
MATERIALS and METHODS spermatophores or spermatangia (sperm vesicles) to deter-
mine if any had mated before collection.
Berryteuthis anonychus was collected as by-catch during a The relationship between latitude and mantle length
United States National Marine Fisheries Service survey of was evaluated using standard regression analysis, and the
salmon in the northeast Pacific (Carlsonet al., 1999). Sam- significance ofthe population regression was tested using
ples were collected during 6 to 17 May 1999 at seven analysis of variance (Zar, 1996:338—343). Normality of
the size-frequency distribution at each station was tested
Please address all correspondencetoJ. R. Bower;e-mail: akai- using normal quantile plots (Sokal & Rohlf. 1995:118—
[email protected] 122). Male and female mantle sizes were compared at
Page 310 The Veliger, Vol. 45, No. 4
each station using the Mann-Whitney test. The relation- data are normally distributed at all other stations except
ship between latitude and the proportion offemales in the #11, where data are negatively skewed due to differences
catch was evaluatedusing Chi-square analysis (Zar, 1996: in size and abundance of males and females.
562-565); the two southernmost stations, where more
than 20% of the samples could not be sexed, were ex-
cluded from this analysis. The relationship between ma- Sex
turity stage (dependent variable) and latitude (indepen-
dent variable) was examined using polytomous logistic
regression. Significance in all tests was accepted at the P Females were both larger and more numerous than
= 0.05 level. males at the northern stations. At each station north of
42°00'N, male and female sizes differed significantly.
RESULTS This size difference was most distinct at Station 11,
mm
where the mean female size was 14 larger than that
Size
of males. The proportion of females in the catches in-
Mantle lengths (ML) ranged from 10.3 to 102.2 mm creased from south to north (Table 2) and followed a sig-
and increased significantly in both sexes from south to nificant linear trend. No females had discharged sper-
north (Figure 2, Table 2). Station #22 was exceptional in matophores or spermatangia present in the buccal mem-
mm
having a wide size range (10.3-75.7 ML), including brane or inner mantle wall, suggesting that none had mat-
eight specimens larger than 64 mm ML. Size-frequency ed before collection.
.,
J. R. Bower et al., 2002 Page 311
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Page 312 The Veliger, Vol. 45, No. 4
165°W
^
49-01 female
Sta. 11
n=30 [~~] male
unident
I I
0^ 145°W
30-
42-07N 47-02N
Sta. 20 Sta. 37
118 n=30
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123456789 123456789
10 10
Mantle length (cm)
Figure 2. Frequency distributions for mantle size ofBenyteuthi.s anonychus collected in the northeast Pacific along 145° and 165°W.
Maturity ranged in size from 64.1 to 94.8 mm ML; 71% were
collected at the two northernmost stations. The seven ma-
Both sexes showed a clear pattern of increasing ma-
ture males collected at and south of41°02'N differed sig-
turity from south to north (Figure 3), and the relationship nificantly in size from those at the two northernmost sta-
between maturity stage and latitude was significant. At tions (Mann-Whitney test, P < 0.001). Of males larger
each station, males were generally in a more advanced than 75.8 mm ML, 94% were mature. Females ranged in
maturity stage than females. Mature (stage V) males oc- size from 23.3 to 102.2 mm ML, but none were mature.
curred at four stations between 40°15' and 49°0rN and Of the most advanced female maturity stage collected
(stage III), 89% were collected at the two northernmost
Table 2 stations. The five stage III females collected at and south
Median mantle length (ML) and % offemales in the catch otfwo42°n0o7r'thNedrinfmfoesrtedsstiagtniiofniscan(tlMyaninn-sWizheitfnreoym ttheosst,e aPt th<e
of Berrxtemhis anonychus at each station. 0.001). Station #22 was again exceptional in being the
Latitude Longitude Median ML only southern station where advanced maturity stages of
(N) (W) Station # mm Female % both sexes were collected.
(
49°or 165° 1 95.7 11 DISCUSSION
47°02' 145° 37 92.2 63
42°07' 165° 20 69.6 38 Berryteiithis anonychus collected during spring in the
41°02' 145° 31 46.5 42 northeast Pacific increased in size and maturity from
40°55' 165° 21 39.3 39 south to north. Didenko (1990, in an abstract from the
40°15' 165° 22 38.1
39°04' 145° 29 26.8 5th All-USSR Conference on Commercial Invertebrates
in 1990) reported similar trends in size and maturity with
- N11
J. R. Bower et al., 2002 Page 313
J
loo-
49-01
female
Sta. 1
se
n(m)=7 male
n(f)=23 145 W
100- 100-
J
o 42-07N 47-02N
c
50- Sta.20 50- $ Sta.37
<D n(m) =64 P n(m) =11
n(f)=45 n(f)=19
J 11
100- 100-
> 40-55N 41-02N
50-i Sta.21 50- Sta.31
0) n(m) =19 n(m)=41
DC n(f)=14 n(f)=27
0-,
I
100 100-
40-15N 39-04N
501 sta.22 50 Sta.29
1 n(m) =1 n(m) =23
n(f)=4 n(f)=27
IV VI IV VI
Maturity stage
Figure 3. Relative frequency distributions for maturity stage ofBerryteiithis anon\chus collected in the northeast Pacific along 145°
and 165°W.
latitude in this area during spring and summer. In the and salmonids at 45°30'-51°N prey heavily on 80-100
absence of known northward currents in this area, the mm ML squid (Pearcy et al., 1988). In the Ridge Domain
simplest explanation for these patterns is that B. anony- and Alaska Stream north of the Subarctic Current, small
chus actively migrates northward during spring and sum- (< 60 mm ML) Gonatus spp. squids replace the larger
mer. B. anonychus as the main cephalopodprey ofboth Pacific
Ommastrephes bartramii, anotherpelagic squid distrib- pomfret and salmonids (Pearcy et al., 1988; Pearcy et al.,
uted widely in the North Pacific (Clarke, 1966), shows a 1993), suggesting that it becomes more difficult to prey
similartrend ofincreasing size with latitude as itmigrates on B. anonychus as it increases in size.
northward during summer and fall. It hatches in subtrop- Males matured at a smaller size than females. These
ical waters, migrates to feeding grounds north of 41°N, data are consistent with those ofother gonatids, including
then returns south of 32°N to spawn (Murata & Naka- Berryteiithis rnagistermagister(Berry, 1913), Gonatopsis
mura, 1998). As it migrates northward along 165°W, borealis Sasaki, 1923. and Gonatus onyx Young, 1972
modal mantle lengths increase from 15 cm near 37°N to (Arkhipkin et al., 1996; Nesis, 1997). The occurrence of
40 cm near 46°N (Murata & Hayase, 1993), which is a small maturing specimens south of 41°N suggests that
mean increase in mantle length of about 19% per degree early maturing forms may occur in southern waters. Ear-
latitude. Berryteiithis anonychus modal mantle lengths in ly- and late-maturing groups have been reported in other
the present study increased from about 3 cm near 39°N pelagic squids, including G. borealis, and the ommastre-
to about 10 cm near 49°N, for a similar mean increase in phids Sthenoteuthis pteropus (Steenstrup, 1855), S. oual-
mantle length of about 23% per degree latitude. aniensis (Lesson, 1830), and Dosidicus gigas (Nesis,
The sizes ofBerryteiithis anonychus found in predator 1997; Masuda et al., 1998).
stomachs collected in the Subarctic Current in the north- Both sexes were of similar size and abundance at
east Pacific during summer roughly correspond with the southern stations, but females became larger and more
relation between body size and latitude seen in the present numerous than males at the noithern stations. Differential
data. Pacific pomfret (Bramajaponica) at 49-52°N prey growth of the sexes is common in cephalopods as they
heavily on > 70-80 mm ML squid (Pearcy et al., 1993), approach maturity (Forsythe & Van Heukelem, 1987),
