Leeches of Laguna Volcán, Bolivia, Including a New Species of Helobdella (Clitellata: Hirudinea)

12 Pages·2001·0.66 MB·English

by  Mark E. Siddall

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3313, 11 pp., 7 figures January 30, 2001
Leeches of Laguna Volca´n, Bolivia, Including
a New Species of Helobdella
(Clitellata: Hirudinea)
MARK E. SIDDALL1
ABSTRACT
Three species of leeches were found in Laguna Volca´n in Departmento de Santa Cruz,
Bolivia. None are known to be sanguivorous. Two of the species found, Semiscolex similis
and Helobdella triserialis, are known to be broadly distributed in South America. A new
species, Helobdella bolivianita, is described. Diagnostic characters for this species include a
nuchalscuteonsomiteVIIIandthepossessionbothofcompactsalivaryglandsandofdiffuse
parenchymal salivary tissue.
INTRODUCTION (Dejoux, 1992), but exclusively from the
Lago Grande on the Peruvian side.
The hirudifauna of Bolivia remains essen-
From September through November of
tially unstudied. Ringuelet (1953) wrote the
1999, the Center for Biodiversity and Con-
only existing account of leeches in Bolivia
servationoftheAmericanMuseumofNatural
based on four specimens mailed to him by
History conducted faunistic surveys in the
Prof. Harry Marcus of the University of Co-
high Andean portion of Bolivia in collabora-
chabamba, Bolivia. Three of the four speci- tion with Colleccio´n Boliviana de Fauna, La
mens were identified to species (Helobdella Paz, and another small excursion for aquatic
duplicata, Helobdella obscura, and Semis- invertebrates in the lowland under theauspic-
colex similis). Later Helobdella titicacensis, es of the Museo de Historia Natural Noel-
originally described from Peru (Ringuelet, Kempff Mercado. This study details findings
1959), and other leeches were listed among fromthelatterandconstitutesonlythesecond
faunistic samples taken from Lake Titicaca account of leeches in Bolivia.
1AssistantCurator,DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory.
2 AMERICAN MUSEUM NOVITATES NO. 3313
Fig. 1. Laguna Volca´n, Departmento Santa Cruz, Bolivia.
MATERIALS AND METHODS ORDERARHYNCHOBDELLIDA
BLANCHARD,1894
Leeches were collected in Laguna Volca´n
SUBORDERHIRUDINIFORMES
(fig. 1), Departmento de Santa Cruz, Bolivia CABALLERO,1952
(18(cid:56)07(cid:57)17(cid:48)S, 63(cid:56)38(cid:57)88(cid:48)W) on November 14,
FAMILYHIRUDINIDAE
1999. Laguna Volca´n is situated in red sand- WHITMAN,1886
stone foothills of the Bolivian Andes (1100
SUBFAMILYSEMISCOLESCINAE
m) and is part of the Rio Grande drainage.
(SCRIBANANDAUTRUM,1934)
Despite its name, it is not a caldera lake and
is not in a geologically active area. Leeches Semiscolex similis (Weyenbergh, 1879)
found attached to submerged stems ofaquat- Figure2
ic plants, and on the underside of submerged
MATERIALEXAMINED:Free-livingfromLa-
branchesalongtheshorelineofthelakewere
guna Volca´n, Departmento de Santa Cruz,
relaxedwiththegradualadditionof50%eth- Bolivia, 18(cid:56)07(cid:57)17(cid:48)S, 63(cid:56)38(cid:57)88(cid:48)W, 14 No-
anol and fixed either in 95% ethanol or in vember 1999, hand collected by M. Siddall
10% formalin. and C. Specht; determination by M. Siddall.
Examination of external morphology and One specimen fixed in 10% formalin, stored
dissections were accomplished with a Nikon in 70% ethanol (AMNH 4220, Annelida);
SMZ-U stereo microscope with a SPOT-RT one specimen fixed in 10% formalin, stored
digital camera. Specimens prepared for his- in 70% ethanol (Museo de Historia Natural
tology were dehydrated through ethanol, Noel-Kempff Mercado, Santa Cruz, Bolivia,
cleared with Hemo-D, and infiltrated with uncataloged); one specimen fixed and stored
Paraplast Plus in a Tissue Tek II tissue pro- in 95% ethanol (AMNH 4221, Annelida);
cessor. Longitudinal sections (5 (cid:109)m) were one specimen fixed and stored in 95% etha-
stained with Haematoxylin and Eosin/Phlox- nol(MuseodeHistoriaNaturalNoel-Kempff
ine and examined with an Olympus BX50 Mercado, Santa Cruz, Bolivia, uncataloged);
compound microscope at low power. one specimen fixed and stored in 95% etha-
2001 SIDDALL: LEECHES OF LAGUNA VOLCA´N, BOLIVIA 3
Fig. 2. Semiscolex similis. (A) Anterior somites with eyes dorsally in II, III, IV, V and VII. (B)
Genital region with six annuli (arrows) ventrally between male and female gonopores. (C) Posterior
somites showing four annuli, three annuli, and one annulus respectively for somites XXV, XXVI, and
XXVII.
nol held at (cid:50)80(cid:56)C (AMNH 100016, Frozen Kempff Mercado, Santa Cruz, Bolivia, un-
Tissue Collection). The separation of male cataloged); one specimen fixed and stored in
andfemalegonoporesbysixannuli,presence 95%ethanolheldat(cid:50)80(cid:56)C(AMNH100020,
of five pairs of eyes (on II, III, IV, V, and Frozen Tissue Collection). The annulation,
VII), somites XXV quadrianulate, XXVI tri- absence of nuchal glands and presence of
anulate, and XXVII uniannulate, all are con- three rows of black tipped papillae are con-
sistent with descriptions for this species sistent with descriptions of this highly vari-
(Weyenbergh, 1879; Cordero, 1937a, 1937b; ablepan-Americanspecies(Blanchard,1849;
Ringuelet, 1944a, 1944b). Blanchard,1896; Weber, 1915; Cordero,
1937a, 1937b; Ringuelet, 1943, 1944a,
ORDERRHYNCHOBDELLIDA 1944b; Klemm, 1982).
BLANCHARD,1894
FAMILYGLOSSIPHONIIDAEVAILLANT,1890 Helobdella bolivianita, new species
Helobdella triserialis (Blanchard, 1849) Figures4–7
Figure3
HOLOTYPE (fig. 4): Free-living from La-
MATERIALEXAMINED:Free-livingfromLa- guna Volca´n, Departmento de Santa Cruz,
guna Volca´n, Departmento de Santa Cruz, Bolivia, 18(cid:56)07(cid:57)17(cid:48)S, 63(cid:56)38(cid:57)88(cid:48)W, 14 No-
Bolivia, 18(cid:56)07(cid:57)17(cid:48)S, 63(cid:56)38(cid:57)88(cid:48)W, 14 No- vember 1999, hand collected M. Siddall and
vember 1999, hand collected by M. Siddall C. Specht; (deposited in Museo de Historia
and C. Specht; determination by M. Siddall; Natural Noel-Kempff Mercado, Santa Cruz,
fourspecimensfixedin10%formalin,stored Bolivia, uncataloged); body length 15.4 mm,
in 70% ethanol (AMNH 4222, Annelida); maximal width 2.6 mm, fixed in 10% for-
five specimens fixed in 10% formalin,stored malin, stored in 70% ethanol.
in 70% ethanol (Museo de Historia Natural PARATYPES: Free-living from Laguna Vol-
Noel-Kempff Mercado, Santa Cruz, Bolivia, ca´n, Departmento de Santa Cruz, Bolivia,
uncataloged);fivespecimensfixedandstored 18(cid:56)07(cid:57)17(cid:48)S, 63(cid:56)38(cid:57)88(cid:48)W, 14 November
in 95% ethanol (AMNH 4223, Annelida); 1999, hand collected M. Siddall and C.
five specimens fixed and stored in 95% eth- Specht. Six mature specimens and 10 im-
anol (Museo de Historia Natural Noel- mature fixed in 10% formalin, stored in70%
4 AMERICAN MUSEUM NOVITATES NO. 3313
and 11 immature specimens fixed in 10%
formalin, stored in 70% ethanol (Museo de
HistoriaNaturalNoel-KempffMercado,San-
ta Cruz, Bolivia, uncataloged); one sexually
mature fixed and stored in 95% ethanol held
at (cid:50)80(cid:56)C (AMNH 100009, Frozen Tissue
Collection);sixmaturespecimensandsixju-
veniles fixed and stored in 95% ethanol
(AMNH 4228); seven mature specimensand
six juveniles fixed and storedin95%ethanol
(Museo de Historia Natural Noel-Kempff
Mercado, Santa Cruz, Bolivia, uncataloged).
ETYMOLOGY: Name refers to the combined
violet and yellow appearance of this leechin
thelivingstate,whichresemblesacomposite
mineral, Bolivianite (or ametrine), a combi-
nationofamethystandcitrineuniquetoeast-
ern Bolivia.
DIAGNOSIS: This species is distinguished
from other scutiferous species of the genus
by having paired salivary glands at the base
of the proboscis as well as diffuse salivary
tissue in the parenchyma, subdivided annuli,
six pairs of gastric caeca including divertic-
ula, and six pairs of testisacs.
FORM(fig.4):Bodylanceolate,broadestin
posterior half; somites I through IV forming
somewhat broadened head region; dorsum
convex,withinconspicuouspapillaeinsome;
venter flat to slightly concave, without pa-
pillae; anterior sucker oval; mouth pore sub-
terminal; caudal sucker circular, concave,di-
rected ventrad, diameter smaller than width
ofposteriorsomites;middorsalnuchalglands
and scute in VIIIa1/a2.
EYES (fig. 4): One pair, punctiform to tri-
angular, at junction of III and IV.
ANNULATION (fig. 4): Somites I and II un-
iannulate; III and IV biannulate; V through
XXIV triannulate each annulus subdivided;
Fig. 3. Helobdella triserialis exhibiting ste- XXV and XXVI biannulate and subdivided,
reotypical three longitudinal rows of black-tipped XVII uniannulate but no distinction from
papillae. caudal sucker middorsally.
COLOR AND PATTERN (fig. 4): When alive,
anterior one-third of body appearing violet
ethanol (AMNH 4224, Annelida); one ma- blending to yellow posteriad; chromato-
ture cleared in Hemo-D (AMNH 4225, An- phores arranged in approximately 30 faint
nelida); one dissected sexually mature, fixed longitudinal arrays dorsally, and approxi-
in 10% formalin, stored in 70% ethanol mately 20 faint longitudinal arrays ventrally;
(AMNH 4226, Annelida);onesectionedsex- dorsally one pair of solid paramedial lines
ually mature, fixed in 10% formalin, mount- from IV to XXIII or XXIV, becoming inter-
edon22glassslides(AMNH4227.1through mittent more posteriorly, a second pair of
4227.22, Annelida); seven sexually mature lines, fainter and more lateral from VII
2001 SIDDALL: LEECHES OF LAGUNA VOLCA´N, BOLIVIA 5
Fig. 4. Holotype of Helobdella bolivianita (A) dorsal view, (B) ventral view, (C) anterior somites
with middorsal nuchal scute on VIII, and (D) midbody somites showing typical dorsal pigmentation.
6 AMERICAN MUSEUM NOVITATES NO. 3313
Fig. 5. Diagram of internal anatomy of Helobdella bolivianita illustrating the relative position and
shape of the proboscis (pr), salivary cells (sc), testisacs(ts),gastriccaeca(gc),intestinalcaeca(ic),and
anus (an).
2001 SIDDALL: LEECHES OF LAGUNA VOLCA´N, BOLIVIA 7
through XXIII, supramarginal lines fainter;
ventrally two pairs of paramedial lines, the
innermost being stronger. Anterior marginof
oral sucker (I, II) considerably less pigment-
ed than midbody somites.
REPRODUCTIVE SYSTEM (figs. 5, 6): Male
and female gonopores separated by one an-
nulus, male at XII a1/a2, female at XII a2/
a3;sixpairsoftestisacsvisiblehistologically
atXIII/XIVthroughXVIII/XIX;spermducts
exit parenchyma in XIII, extend posteriad to
XVI, and fold back to XV or XIV such that
if unfolded would reach XVI before return-
inganteriad,spermductsemptyintoatriaan-
terioventrally without preatrial loops; atria
piriform approximately at 45(cid:56) to midline;
ovisacs robust but simple, extending to XV.
ALIMENTARY TRACT (figs. 5, 7): Proboscis
slightly thicker at base than tip, in membra-
nous sheath, base of proboscis at XII in re-
laxed state; salivary cells arranged both as a
pair of glandular masses at base of proboscis
and diffusely in parenchyma, ductules of the
latter forming a bundle inserting into thefor-
mer, oesophagus simple, not recurved; gas-
tric chambers with digitiform caeca, six in-
cluding postcaeca (diverticula), first five in
XIV through XVII, postcaeca from XIX
through XXIII or XXIV; intestinefromXIX/
XX,fourlobesbutnotpronouncedlycaecate;
anus at XXVI/XXVII.
REMARKS: No other scutiferous species of
leech is known to possess both compact sal-
ivary glands and diffuse salivary tissue. The
presence of a nuchal scute on VIII is a clear
synapomorphy (Light and Siddall, 1999) for
a subset of species in the genus Helobdella,
indicating that H. bolivianita is allied with
the type species of the genus, Helobdella
stagnalis (L.). In South Americathereare11
known scutiferous species, most of which
can be readily distinguished from H. boli-
vianita. Although Adaetobdella xenoica
(Ringuelet,1975)Sawyer,1986,hascompact
salivary glands, six pairs of testisacs and a
Fig. 6. Median reproductive anatomy of He- scute on VIII, it has seven crop chambers, a
lobdellabolivianitarelativetosomaticgangliaXI preatrial loop for each sperm duct, and no
throughXVI.Abbreviations:at,atrium;sd,sperm posterior crop diverticula and does not ex-
duct; ov, ovary.
hibit subdivided annuli (Ringuelet, 1975,
1978b). The most broadly distributedspecies
of scutiferous leech in South America is He-
lobdella scutifera Blanchard, 1900, known
from as far south as Tierra del Fuego
8 AMERICAN MUSEUM NOVITATES NO. 3313
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2001 SIDDALL: LEECHES OF LAGUNA VOLCA´N, BOLIVIA 9
(Moore, 1911) and as far North as Rio specimen attributed by him to H. duplicata
Grande do Norte, Brazil (Cordero, 1937a). details only external morphology and does
Blanchard (1900) noted dorsal paramedial not notethestronglymetamericpigmentpat-
linesinsomeindividualsofH.scutiferarem- tern otherwise characteristic of this species
iniscent of Glossiphonia complanata and su- (Moore, 1911). In later work (Ringuelet,
perficially similar to those described herefor 1944b)thecharacteristicsattributedtoH.du-
H.bolivianita.However,H.scutiferahassal- plicata, ‘‘Color liso, con los annilos a3 ma´s
ivary cells diffusely arranged in the paren- pigmentados, o liso con 2 estr´ıas dorso-lon-
chyma, a very long proboscis extending to gitudinales’’ were not completely in accord
and folding back anteriorly at XIV, ovaries with Moore’s (1911) description of H. dupli-
only as long as XIV, and sperm ducts that cata. Moore (1911) noted two broad and
reach XIX (Blanchard, 1900; Weber, 1915; faint paramedial bands of pigmentation but
Ringuelet, 1978a). Helobdella diploidesRin- at six muscle bundles wide they were in no
guelet, 1948, although it lacks any observ- sense distinct longitudinal lines on the dor-
able pigmentation, exhibits subdivided an- sum like those of H. bolivianita. Eventually
nuli from VII through XXV and a scute in Ringuelet (1985) appears only to have re-
VIII similar to H. bolivianita. Otherwiseitis quired subdivided annuli and a scute for in-
unique in having exceedingly short sperm clusion in H. duplicata, thus obviating the
ducts, recurving anteriorly at XIV, and atria need for dissection. As such, it is quite pos-
that are distinctly at right angles to the mid- sible that the one specimen from Cochabam-
line (Ringuelet, 1948). Helobdella godeti ba, Bolivia, was not H. duplicata. Whether
Weber, 1916, Helobdella simplex Moore, or not it was Helobdella bolivianita cannot
1911, and Helobdella montevidensis Corde- bedeterminedfromtheinformationprovided
ro, 1937, though scutiferous, do not have (Ringuelet,1953).ThepresenceinBoliviaof
subdivided annuli and each has obvious dor- the other two species noted here is unre-
sal papillation. The most similar species and markable. One individual of Semiscolexsim-
one that individuals of H. bolivianita may ilis also was among the leeches detailed by
previously have been confused with is He- Ringuelet(1953)fromCochabamba,andHe-
lobdella duplicata Moore, 1911, originally lobdella triserialis is found from Concep-
described from Patagonia but now attributed cio´n, Chile (Weber, 1915) through Northern
to a broader South American distribution. Michigan, USA (Klemm, 1982).
Helobdella duplicata has subdivided annuli, The presence both of compact salivary
sixpairsofgastriccaecaincludingpostcaeca, glands at the base of the proboscis and of
six pairs of testisacs, a scute in VIII, and a diffuseparenchymalsalivarytissueinHelob-
caudal sucker directed strongly ventrad della bolivianita is unprecedented in Hiru-
(Moore, 1911). However, numerous features dinea. Two principlecladesofleechesexhib-
distinguish H. duplicata from H. bolivianita, it the derived condition of compact salivary
most notably a lack of compact salivary glands (Light and Siddall, 1999): a mono-
glands, a transverse metameric pigmentation phyletic subset of Placobdella species and
on annulus a1 of each somite, globular atria thegenusHaementeria.Althoughnotinclud-
with anteriodorsal insertion of spermducts, edincurrentphylogeneticassessmentsofthe
ovaries that reach XVII, and the presence of Glossiphoniidae (see Light and Siddall,
pronounced lateral caeca of the intestine 1999), the genus Adaetobdella Ringuelet,
(Moore, 1911). 1978 comprises species previously in thege-
nus Helobdella but which lack parenchymal
DISCUSSION salivary tissue and possess a pair of compact
salivary glands set-off from the base of the
Ringuelet’s (1953) note, the only preexist- proboscis by their bundled salivary ductules
ing taxonomic account of leeches from Bo- (Ringuelet, 1978b). Because the genus Hae-
livia, recorded the existence of Helobdella menteria currently is supported as sister tax-
duplicata, Helobdella obscura and an undes- on to Helobdella, it may have been reason-
cribed Helobdella species in samples sent abletoconcludethatthepresenceofcompact
from Cochabamba. His description of the salivary glands is a synapomorphy for the
10 AMERICAN MUSEUM NOVITATES NO. 3313
generaHaementeriaandAdaetobdella.How- 307–310. Dordrecht, The Netherlands:
ever, the existence of both kinds of salivary Kluwer.
tissue in the scutiferous H. bolivianita, and Klemm, D. J.
only compact tissue in another scutiferous 1982. The leeches (Annelida: Hirudinea) of
North America. Aquatic Biology Sec-
species, Adaetobdella xenoica, puts those
tion, Environmental Monitoring and
characters in conflict. Because the type spe-
Support Laboratory, Office of Re-
cies of the genus, Helobdella stagnalis, has
search and Development, U.S. Envi-
a scute, retaining scutiferous species in the
ronmental Protection Agency. Cinn.,
genus Helobdella would seem to be prudent. Ohio 45268.
Light, J. E., and M. E. Siddall
ACKNOWLEDGMENTS 1999. Phylogeny of the leech family Glossi-
phoniidae based on mitochondrialgene
I thank the Center for Biodiversity and
sequences and morphological data. J.
Conservation of the American Museum of Parasitol. 85: 813–823.
Natural Historyforfundingandcoordinating Moore, J. P.
this expedition. The collegiality and gener- 1911. Hirudinea of Southern Patagonia. Rep.
osity of Maria-Esther Montan˜o of the Museo Princeton Univ. Exped. Patagonia,
de Historia Natural Noel-Kempff Mercado, 1896–1899. 3: 669–689.
Santa Cruz, Bolivia facilitated collectionand Ringuelet, R. A.
exitofspecimens.IthankChelseaSiddallfor 1943. Sobre la morpholog´ıa y variabilidadde
Helobdella triserialis (Em. Bl.) (Hiru-
her expertise as field guide and chauffeur as
dinea,Glossiphoniidae).NotasMus.La
well as for her assistance with thecollecting.
Plata 8: 215–240.
Paula Mikkelsen kindly made available re-
1944a. Synopsis sistema´tica y zoogogra´fica de
sources and knowledge in her histology lab-
los hirud´ıneos de la Argentina, Brasil,
oratory.IthankMarieLawrenceforherthor-
Chile, Paraguay y Uruguay. Rev. Mus.
ough histological and curatorial work, as
La Plata 3: 163–232.
well as Steve Thurston for skillfully illus- 1944b. Los hirud´ıneos Argentinos de los ge´-
trating the internal anatomy of the new spe- neros Helobdella R. Bl., Batracobdella
cies. Vig., Cylicobdella Gr. y Semiscolex
Kinb. Rev. Mus. La Plata 4: 5–94.
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