PICTORIAL KEYS TO THE AQUATIC MOLLUSKS OF THE UPPER SUSQUEHANNA by Willard N. Harman Biological Field Station Cooperstown, N.Y. Occasional paper No. 9 Januarv 1982 Biology Department State University College at Oneonta II
PICTORIAL KEYS TO THE AQUATIC MOLLUSKS OF THE
UPPER SUSQUEHANNA
by
Willard N. Harman
Biological Field Station Cooperstown, N.Y.
Occasional paper No. 9
Januarv 1982
Biology Department State University
College at Oneonta
II
This paper hM be~:u, '(T:.;;,.t"~E;r.:. :eOl:" th<i:0 ,-'>Be of l~1J::en a.nd students interested in the natural h:tst;)r:r 0: "~he ~e"c of Hew York State.
It incluc"lesTli?",-;:,;:YJ.""l C:Gi:::C:('.;3:'E'l],;lg; t.hG 1c;:;?~:.
s:-;:ecJ,<;;'i',:n:i;Ui" O:f:7).n
D~d ecological aspects of the biology of au,x fl'eshwater mussels, as well as pictorial l:ti'it cd.'the species collected in the Cou:o:tJ ~s pre~ented in Table 1. Also included ar,~ -c,:b.c'" or Irmd snails (Table 2.) found i.n 'the Lake.: (:,,:n'iT.,Lr,;il:l::;? for th05c interested in the non-aquatic molluscEL,
The f're5n\;atE"Y fi' F~:U.i3
Gastropoda.. In COX:;::;,::::'",!,?';':" .;~;:;
Gastropoda have encapsulated '[;&;0 They comprise t;f.f'"
Mollusca of the Class '::1..:1.8 class, all freshwater tns production of yolky,
18~d (Hunter, 1964). '3_:~~ K"::'i;Go1:n,"sr!.chia and the
Pulmonata.
The P'L>08(}b:F."'J:1:~
Yi opercU:L:,~
lit,'co:,"fJ, he.v~
Ii1"=
PJter
can affecte~
;,~:n(j,
lliB2es F:"",:;;.:C'.i2ti;~· prosclhraD.cus
::~L1:::;-::,I;:; f; and these t,;o groups ..',', g,nees'{;:nal stock
~~;L?S'2: sLd.ls ht'HI toi.n 1.0\-[
e:'.~':",ob~~£;..nchia tJ.C quired adaptations-.", ·:;;~t:.~,'·~·:;_r::~u.{:;
"":,k.:C r~a;,:'':.lE.:
'~he f;\lb3'.:,};'''Ste. ~:'4"~d.~~Q \U_Lb_
~.t'.
the families Valvatidae ~ aTe characterized by is s. rigid or leathery disc prot(~cting t.he snail whenever it is in central New York, except the Valvati&~e being easily recognized because of the are closely allied to marine" presUllJAbl;y (Hunter, 1952) • To develop an osmoregulatoj:-;:/ salt coneentrations. for life in ha'v'e developed gills tha.t C:8.vities and thus are less Some members of the Vivips.J:':LdE£ fT"~~ans-M~u, v19~6) •
This avoids th:":'?"l':", 'hhUe plowing through the substrate They are also better hidden i,;.t8.t ionar}" posit ions.
St,,,;{lo!!rmatophora and Basommatophora, r·lelD~0·:S:::s
()X"{i,C;,I'2
& 'c,'&.il:',L,,,';'. ~~,,->:(L.'i.
o? tn.~ tervl..:t~z·i~rial sna.ils and
slugs, have tvo
The Pulmonata &~'~
'OOTne <.d:, the tips of tbe ~longer pair. l',:~ __ , ,~",,_ O~,'>(i~~r :Baf:::_~~\rf:11L~tophora
They have onl;y" OUE; (};..c' ·i~8:D.t8..~~t;::-;8 ':iJ',l:.!, :,il'';'' '::';:F;;8 are v,at on them but near the bases (see coy(;?~" :L1J_;':(~'C:;;'B,tj,()Z;), LFJ':di f!ilL.::.13 i2~nd their empty shells frequently are i'm.met ;ii::.1) ";;t:L2:,o to ident:Lfy- a.quetic
nE1de by stud~utst.bE:!Jl tG
'I#he
';7[1,t",: c"
,u1: ,::l1811 the ';;0 02' 8'~
x·e8~)-=.
ZJ~l'1d],s
snails should keep them in m,:u:m. who have tried to use these keys b.z;;\TS J.&.l2d snails I If the soft tion of tbe eyes can be deterr:c0:u,;;cL couplets may be those of If:lx:dL spire raised only slightly abcnr~e tl1.e species of Valvata (greatest dia:mett':;;Y'
III
The basommatopho!'aTI families in central New York (Lymnaeidae, Physidae, Planorbid.a.e, Wld f.l.L1cyLi.c...a.e) are characterized by combined sexes (monoecious species), lack of c>:p.eZ-CUl8~l' no tru.e gills, and a. highly vascular mantle cavity that is used as a l1mg, OJ..tyge!:i is obtained from either air or water, which enters the ma.Tltle cav:it.y th:!.~ougb an opening known as the pneumostome. The Planorbidae and Anc:rl:l:.dae have developed secondary gills from folds of tissue associated with. but oriented outside, the mantle cavity.
The freshvater J:nl]~IT£);:la,tes ay-parently evolved from ancestors that were air-breathing land snails (Bu.I."1ter, 1952). 'J'hese families illustrate different degrees of ada~tation to aqL~tic life, which may represent steps in a process of evolution. Some 3,UtJ10X':E have consid.ered the am;phibious members more primitive than the fm:Lll..8 that nev'':er use atmospheric air (Morton. 1955). But, each family has become ,.;ell "3,dB,:9t;(~dto the environment in which it lives.
The members of the LjI'1:imaeidae (illustrated on the front cover) depend for respiration entirely on the rich vascular tissues of their otherwise simple mantle c~.vities. 'l11:L, lJlung" 18 f'unctional whether filled with air or vater (Hunter~ 1953b)0
The ma.:ntle ca:v:tt:ies oj: t.he Pbssidae are similar to those of the Lymnaeidae. In addition, the mantle possesses digitate outgrowths that lie over the outside of the shell and increase stD~!ace/volume relationships of the snails. theoreti cally resulting in less dependence on atmospheric air. Hovever, many species of Physidae seem j1..lSt as inclined as the Ii;rmnaeidae to remain at the surface, where they can takt~ ,:LiZ' :tnto the mantle ca'fity"
Some species in the fa""ily Planorbidae have developed a lobe(s) of tissue that lies outside the mantle cavity and serves as a gill. This advanced modification gree.tl~r :tnrpr07es their capability of extracting oxygen from water. Few membe!:";.', ci': the' vclunta,rily el11erge from the vater. but they often remain (:;'f)!:E; to the" surface and take air into the mantle cavity.
The .Ancylidae c m: :f::c'.':~,~:·:c.:t,e)· limpets" have gills much like those of the planorbid snails ,. 'I":,),,!:'.;' T'-:C 'tJ.2:"cshave become so efficient that large mantle cavities are ~hose of the Fncylidae have become so reduced that they to satis~~r respiratorj needs. Limpets are normally fmu:,d (')2 h'.::,', :5.':;.::"'\:<f1''':'e substr&tes below tbe vater surface.
The me8.L'1 ratic of sbell \;!(dgh-t, to llodJr 'sleight is appreciably less in the Pulmonata thr;tTI iD 2~·':;sobr(':xl(.;hi' The rels.tively light shells of the Pulmonata. '.re"ce esse;:;tial iL \:h0 deveJo:;;.r.ment of their common habit of grazing upside dOT;IT; c::m the f"urf'a,C€ films of q,uiet waters. In general the Pulmonata are mO!t'; tlct i ye ':ll&n tn,,; Prosobrw"1chia. moving over vegetation. substrate~ and slITf~ce fiJ~s at rates of up to 0.5 em/sec. Because they are not dependent on 1:n."es"tbj.ug, the:r tolerate exposure to air better than do most prosobranchs BtLried in mud and organic substrates. many pulmonates0
can withstand drJ c:0'J/:.It5.oJ':.8 OtT'ar the greater pErt of the year.
The three factors discussed at-ove - light. weight. rapidity of movement. and ability to ,.rithstanQ expor;n.:tre to air - reB111t in high dispersal rates of the Pulmonata as COllipllL'ed \iith those of' the heavy. more ponderous. water dependent Prosobranchia. Not only are the Pulmonata transported passively by birds, insects ~ and other &.nim.."!.ls, but they disperse actively across marshes and other ephemeral B~U2~ic biotopes. Their ability to move through
IV
marshes provides a particularly important means of dispersal in central New York, where marshes in the litm:ough valleys" serve as portals for migrat ion from one major river system t;o ano\--'her.
'l'HE FRESHWATER CLAMS A.."fill MUSSELS
The fre8h~vater clams ere Mollusca possessing t·'I{i.) shells (Class £ivB.1via) . All local fresh-water bi"lalYes exhibit e);:"tex'uM fertilization of eggs while stored in pouches "."ithin cheir g:i-lls" 'l'hey comprise '1;/",1.) f'amilies ~ the large Unionidae or pearly f'resh-lii?;,t(~:,' lID)j3E,"",ls ~ ,,,2:it! the rn,Jc:u sm~~11e};~ ind:l.viduals ,,vithin the fwnily Sphaeriidae, the or.' pea, clw.mL
The Unioni,dae £J:'e characteri.zed by their large size (up to 15 cm in length) and the fact that during their juvenile st~ses they are parasitic on fishes. Their life cycles are as fOLlows, E&ss dev010ping in the reproductive organs of the females moVe':: rTf)';!" those striJctt1.:;-:,,~~!S U;lto the ll:lffi.'ltle ca,yity. hence to marsupia wi.tbi.rJ th:: ,s2;',;',:;,r'd,liz,ed. s}Jer:w brought into the mantle canty during thE; 3 ri~):-;511'1J. g,ct :br:L There they develop into two-valved larvae ;2~houi; 1. to ~: Ire::!. :L.1 :leug'l;f\ c~,'lle(1, glochidia. The glochidia are released into the ~~ter by the females either passively or actively. In the first instance glochidia are simply spew"d out where they remain suspended near the substrate by virtue of their p:;~acticall;,r neutraI huoye.J:lcy. Any fishes swimming in the area, taking in wat",,!, for :respi1:"ation, ine.c:,"~~rtently pick up the larvae and pass them by their gill filaments. t~n contact the glochidia clamp their valves upon the gills of the fish and assume a parasitic existence. In the second situation portions of the mantle of the female mussel are waved about imitating a small fish (inside back cover). ween a l!~ger fish attacks the clam forcefully expels glochidia in the predator'S face 9 assuring the future of her young. After a period of development that ',1'8,l'ies consider8.bl~y the glochidia drop from the gills. fall to the bottom and take up a free-living existence as adults (Figure 1).
As adults. vater contlll1..ll'l.l.J.y taken into the mantle cavity for respiration and removal of wa.stes brings in e~gae. bacteria and other edible microscopic particles which are filtered on the gills and trans~~rted to the oral region for food..
Members of the rEJfl~ C'::d,~:'llaae are no~ parasitic on fish. the young developing vi.th:tn the \;;c,'t.:,:L the a6.ult stage :i.e reached. Immediately upon release from the fenw~_e "hey 8,:'\~(~ able to Hve independently as filter feeders just as adult m"lJ.oni.dEL
Both families c{ b:Lval·;I,c;;: 32'e ~ :c'cr theiy respective sizes. rather ponderous and water dependent. 'l'herefore the ecolog'{ is similar to the prosobranch snails. Importantly. the UnloLifia>e mUfri.:. I)€; in 'd'8.tcr' 'Where their host fishes occur or they cannot complete their life CyCles. This phenomenon normally restricts them to larger lakes and st.r:e2"ITcS 0
In geologica.l 't.e:..,':m3, fresh water',,;; 2..1"2'[':::('Y short-livedo Therefore. populations of fresh-w"ateY' molJ.u'sks ral',,~ly remain isolated long enough to become fully speciated. even though tempor~{ isolation is ~ornmonplace. For this rea8on~ there are relatively fev spectes of t'r(",sh.-'~i'"ater sne.i.ls and c18ill.S $ but most species are widespread and quite v8.1:'ia"bl.::" '1'0 Sl.lrVl"\re, the;;.' :must '[,clerate such conditions as extremes of temperature, dissolved oAfgen,aJ~calinity, £~d pH. Adaptive plasticity, the ability of evolutionf'-ry lines t.o c'bange't,,11eY,:: e.,.-q:'Osed to various stresses. appears to be extremely important in fresh-water gastropods (Hunter. 1964) and bivalves and to be of selectiv-E: e,(lvantage in eV".'llution. Specializations that render eo species more efficient in one envirorrrr~nt but l~ss capable of colonizing different biotopes are selected against (If'lbel1cic$., 2_962),
TABLE 1. THE AQUATIC MOLLUSCA OF THE UPPER SUSQUEHANNA.
Class GASTROPODA Family Hydrobiidae Subclass Pulmonata Amnicola limosa Order Basomattophora Amnicola lustrica Family Lymnaeidae Amnicola integra
Lymnaea humilis LYmnaea palustris Family Pleuroceridae Lymnaea ~mers.inata Spirodon carinata Lymnaea columella Goniobasis virginica
Family Planorbidae Family Valvatidae Helisoma trivolvis Valvata tricarinata Eelisoma anceps Valvata _sincera Helisoma campanulata Gyraulus 12arv-~s CLASS BIVALVIA Promenetus exacuous Family Unionidae
Lampsilis radiata Family Physidae Lampsilis cariosa
Pbysa heterostrop~ Elliptio complanata Aplexa bypnorurn. Anodonta cataracta Physa integra Anodontoides ferrussacianus Pbysa. gyrina Strophitus undulatus
Alasmidonta undulatus Family Ancylids.e Alasmidonta undulata
Ferrissia parallels. Lasmigona compressa Ferrissia rivularis Alasmidonta marginata
Actinonais carinata Order Prosobranchia Family Viviparidae Family Spbaeriidae
Viviparus georgianus Pisidium casertanum Cipangopaludina chinensis Pisidium compressum
PisidiuIll subtruncatum Sphaerium sulcatum
TABLE 2. TIlE LAND SNAILS OF 'fH:E UPPER SUSQUEH.A..1IfNA (from r~cNa.mara and Harman. 1975)
Order Stylomattophora Ca.rychi1L~ exif£l1u.m Stria.turu milium Gastrocopta pentodo? Striatura exigua Vertigo bolleslana StenotremF~ fraternum Succinea ovalis Triodopsis dentifera Ce.tinella vermeta Triodopsis denotata Anguispira alternata Triodopsis tridentata Discus catskillensis Mesodon oayanus Discus cronkhitei ~iodopBis albolabris Discus patulus . Arion rase iatus Pulletum minuti 5 siJlnml Deroceras la-eve Nesovitrea binneyana Deroceras cf. agreste Mesom,phix inornatus Philomycus flexuolaris Euconulus chersinus p?lygyratus Philomycus togatus Ventridens intertextus Pallifera dorsalis Zan!toides arboreus
VI
4.J
SPERM EJECTED INTO WATER
/''- ' -= .=~'<tW~~~ ,,'"
EGGS GROW t>
TO GlOCHiDlA ~ '"' ~ AND ARE _~"" EJECTED FROM EXHAlHJT SIPHON
\ \
©(; ~~ .~ ~ 3. - fd";~~ \ ,
SPERM T t~KEf\,! .. ('j ~, t~~~~~~~ I~J INHAl Ei\n-S5PHOI'J. ?ASSEJ THr~GUGi-'1 GfLLS, fERT!UnNG THE EGGS
UNfERTILIZED EGGS IN GilLS
METAMORPHOSIS TAKES PLACE. YOUNG CLAMS BURROW OUT OF Gill, FAll TO BOTTOM. AND GROW TO ADUl TS
ADULTS
"
@~ o //
~-U--'<
HOST TISSUES ENCOMPASS PARASITIC GLOCHIDIA WHILE THEY DEVELOP
WHEN PASSING 6. OUT THROUGH
GilLS GlOCHIDIA CLOSE ON FILAMENTS
Figure 1. The life cycle of a :fresh~we.t,,~.r cl:~~ Family Unionidae.
VII
REFERENCES
Buckley, D. A. 1977. The distribution and ecology of the aquatic molluscan fauna of the Black River drainage basin in Northern New York. Occas. Pap. No.6. SUNY Oneonta Bio. Fld. Sta., SUlfY Oneonta.
Burch, J. B. 1975. Freshwater sphaeriacean clams (Mollusca:Pe1ecypoda) of North America. Museum and Department of Zoology. The University of Michigan. Ann Arbor. 96 pp.
Harman, W. N. 1980. Map of otsego Lake In MacWatters, R. C. The fishes of otsego Lake. Occas. Pap. lio. 7. SUNY Oneonta Bio. F1d. sta., SUliY Oneonta.
Harman, W. N. and C. O. Berg. 1971. The freshvater snails of centr&1 New York with illustrated keys to the Genera and species. Search: Cornell Univ. Agr. Exp. Sta•• ENT. Ithaca. 1(4):1:63.
Hubendick, B. 1962. Aspects on the diversity of the freshwater fauna. Oikos. 13:249-261.
Hunter, W. R. 1952. The adaptations of freshwater gastropod,a. Glasgow Nat. 16:34-35.
Hunter, W. R. 1953. The condition of the mantle cavity in two pulmonate snails living in Loch Lomond. Roy. Soc. Edinburg. !Toe. a65:143-165.
Hunter, W. R. 1964. Physio10gic&1 aspects of ecology in non-marine mollusks, p. 83-ll6. In Physiology of Mollusca, vol. 1, by K. M. Wilbur and C. M. Yonge-red.) Academic Press, Nev York. 473 pp.
Jorgenson, C. B. 1966. Biology of suspension feeding. Pergamon, Oxford. 357 pp.
MacNamara, M. C. and W. N. Harman. 1975. Further studies of the Mollusca of the Otsego Lake area. Nautilus, 89:87-90.
Morton, J. E. 1955. The evolution of the Ellobiidae with a discussion on the origin of the Pulmonata. Zool. Soc. London. Proc. 125: 127-168.
Thornton, S. A. 1980. Bathymetric map of Goodyear Lake, p. 716. In 12th Ann. Rept. (1979). SUNY Oneonta Bio. F1d. Sta., SUNY Oneont~
Weir, G. P. and W. N. Harman. 1975. Bathymetric map of Canadarago Lake. otsego County Conservation Assn., Cooperstown.
VIII
(clams)
whorls in more than one plane
(p. 3)
shell coiled
A PICTORIAL KEY TO THE AQUATIC MOLLUSCA OF THE UPPER SUSQUEHANNA WATERSHED
PHYLUM MOLLUSCA
sides of shell parallel, sides of shell not found aD vegetation in parallel, found on rocks slow moving water in rapidly flowing streams
Ferrissia paralella Ferrissia rivularis
:::lass GASTROPODA (snails)
I shell cone shaped
I Family Ancylidae
(the limpets)
-----------~ Class BIVALVIA (p.8;
whorls in one plene
Family Planorbidae (the rams horn snails)
(p. 2)
1
-----
. .
/anceps
Family Planorbidae (from p. 1)
shell large shell small
/at least 7 mm in diameter ~
extremely not extremely flattened flattened
Helisoma I
whorls flattened whorls r~ded side
on on both sides at least one
H.
Promenetus exacuous
~."..
whorls flattened whorls rounded on one side on both sides
less than 2 mm in dia./ I H. trivolvis H. campanulata Menetus dilatatus
greater than 2 mm in dia.
I Gyraulus parvus
2
lack sculpture
whorls on more than one plane (from p. 1)
~ ~ whorls sinistral vborls dextral
~ ~
/ not operculate operculate (having a
trap door covering the
aperture) (p. 5)
Family Physidae (the ear or tadpole snails)
/shell elongate, oily in appearance Aplexa hypnorum Family Lymnaeidae (p. 4)
shell inflated, not oily in appearance
(the pond snails)
Genus Pbysa
/- spiral sculpture
.____--------1 shell e~emely shell moderately Physa heterostropha
inflated inflated
Physa sayii Physa integra
3
Family Lymnaeidae (from p. 3)
(the pond snails)
~ shell thin. fragile
Lymnaea columella
shell thick. resistant
large"adults small. adults 15 mm or more less than 15 mm in height in height
Lymnaea hUDdHs
shell inflated shell elongated Lymnaea emarginata yY!naea palustris
4
shells operculate (trom p. 3)
~Pro~ spire only slightly spire moderately to
elejted extr/emelyele~d
Family Valvatidae '"
whorls inflated whorls flattened
shells carinate Valvata tricarinata
shells without carina. Valvata sincera
Family Pleuroceridae (p. 6)
adults greater than adults less than 15 mm in height 15 DUll in height
Family Viviparidae (p. 6) Family HydrobiidAe (p. 7) (apple snails)
5
Family Viviparidae (from p. 5)
\ shell with shell vithout 4 brown bands vithout bands-
---- shell shouldered Viviparis georgianus Campeloma decise.
shell obese
Cip&Il§0pal.udine. chinensis
Family Pleuroceride.e (from p. 5)
shell globose shell elevated
Spirodon carinate. Goniobasis virginica
6
Family Hydrobiid&e (from p. 5)
shell truncate shell conic or attenuate
I Amnicola limosa
shell conic shell attenuate
Amnicola integra Amnicola lustrica
~
,I
I
I
7
CLASS BIVALVIA (from p. 1)
small, less than 10 mm in length
(clams) ___________
large, 15 to more than 100 mm in length
Family Sphaeriidae Family Unionidae (p. 9)
(the pea or finger-nail clams) (the pearly fresh-water mussels)
beaks posterior beaks anterior or subcentral animal less than 4 mm in length adults greater than 4 mm
in length
Genus Pisidium (pe,elams) Genus Sphaeridium (finger-nail clams)
The species in these two genera are very difficult to identify and are therefore omitted in this work designed for the layman.
8
Fr:..mily UNIONIDAE
(the pearly freshwater mussels) (from p. 8 )
_•.,sJ.p
~""P? ~
hinge with articulating hinge otherwise pseudocardinal s.nd lateral (p. 12) hinge teeth present
shell without rays~ jlhell with rays one of our most COIDID.on (p. 9)
mussels
Elliptio complanata
9
Shell with Rays (from p. 9)
---shell trapazoidal~ shell ovoid found on.ly in Canadarago
Lake
shell with very few or broad rays~
found only in rivers
shell possessing many thin ra:ys, commonly found in lakes and rivers
Lampsilis radiata
/i
shell large, yellow with only a fev thin rays (p. 11)
broad rays. distinct growth linesj background color greenish-yellow
rare.. in rivers
Actininajs carinata
~o
Shell Large, Yellow, With a Few Weak Rays, Only Found in Large Rivers
(from p. 10)
Lampsilis cariosa
11
Hinge Without Articulating Lateral and Pseudocardinal Hinge Teeth
(from p. 9)
~ hinge 'Without teeth hinge 'With teeth
(p. 13)
shell small (never exceeding 8 cm shell often large, common on in length) found only at muddy bottoms everywhere the north end of Otsego Lake
Ar.odontoides ferrussacianus
Anodonta cataracta
12
Hinge With Teeth
shell corrugated OD
posterior s~ope,
found only 1D
rivers
shell
inside of shell yellow-pi~, • pseudocard1n8..l. small and smooth
inside of. shell reddish-Plnk, pseudocardinal large and rough
Alasmidonta marginata
Strophitus undulatus
Alasmidoota undulata
13
I iI-I
ACKNOWLEDGEMENTS
The illustrations contained within this MS are, of course, the single most important part of the work. Although the author is responsible for the front cover and all figures of snails others should be credited as follows:
Ms. Karen Elting, Oneonta, N.Y. Family Unionidae
Heard and Burch (1975) Genus Pisidium Genus Sphaerium