Bijdragen tot de Dierkunde, 50 (1): 235-242 — 1980

Freshwater from Colombia.

A revision of Fuhrmann’s types

by

Ian R. Ball

Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands

Abstract in mis- tant , are occasionally

The type material of Colombian species of freshwater triclads interpreted because the particular specimen is (Fuhrmann, 1914) is revised and the original descriptions curved, or for some other reason the sections are corrected and amplified. Planaria polyorchis is conspecific In Fuhrmann's Planaria not true. other descriptions, with festai (Borelli). longistriata is a respects

typical species of the Dugesia (Girardia) tigrina originally in French, are generally reliable and

group, a group widespread in North and South America. here in are repeated only summary. Planaria paramensis and P. cameliae belong to a group of

Dugesia species characterized particularly by their dorsal

and number of and around testes, exemplified by a species in SPECIES ACCOUNTS

the Caribbean region. Dugesia (Girardia) longistriata (Fuhrmann,

1, INTRODUCTION 1914) (Figs. 2)

work on often is Revisionary aquatic planarians Planaria longistriata Fuhrmann, 1914: 796

hampered by the inadequacies, according to modern Euplanaria longistriata: Kenk, 1930: 292. Dugesia longistriata: Kenk, 1941: 7. criteria, of early species accounts and by the general

of material. It is unavailability type important, Type material. — British Museum (Natural History),

therefore, when types are rediscovered, that they be 1928.1.5.16-18 and 1928.1.5.81-86, alcohol specimens (para- types). BMNH 1928.1.4.245-257, slides as follows: 245-250 reassessed and their status and location docu- wholemounts; 251 oblique sagittal sections; 252-253 sagittal mented. In the I the present paper report on type sections, copulatory apparatus not in the series; 254 sagittal

material of freshwater planarians described from sections (cf. Fuhrmann, 1914, fig. 4); 255-257 frontal sec- tions (cf. Fuhrmann, 1914, fig. 5). All the available slides Colombia, South America, by Fuhrmann (1914), been but have re-examined the new description and figures

which are now housed in the collections of types are based primarily upon the wholemount 245, sagittal series the British Museum (Natural History). Our total 254, and frontal series 255-257. Slide 254 is difficult to interpret because it has at some time been damaged and knowledge concerning the Colombian paludicolan repaired. fauna is derived only from Fuhrmann's publi-

cation and in view of the of interest — resurgence Redescription. The external features

in the of northern Neotropica (Ball, are as described by Fuhrmann (1914; 796) and

Mitchell redrawn here 1971; & Kawakatsu, 1972; Codreanu specimen 245 has been as fig. 1.

& in it to Balcesco, 1973; Gourbault, 1979, press) Living specimens are up 18 mm long, very

is unfortunate that Fuhrmann's actual and of ochre colour with dark specimens agile, a light many

should have remained unknown and unstudied for brown stripes. The ventral surface is pale and lacks

The is in so long. stripes. pharynx pigmented, as other

the that World Accepting principle our task is not to New Dugesiidae, the pigment lying imme-

know beneath name species but to them I am presenting diately the outer musculature of the here corrections and amplifications to Fuhrmann's pharynx.

of the Colombian to The testes are and descriptions species so as numerous, small, occur in

facilitate further monographic work on the Duge- clusters ventrally situated throughout the body- siidae. The original accounts lack histological length. There are three or four testicular follicles information and the the which latter copulatory organs, so impor- anterior to ovaries, occur ventrally 236 I. R. BALL - FUHRMANN'S COLOMBIAN PLANARIANS

found in between the third and fourth intestinal diverticula. Distribution. — This species was

The vitellaria are found throughout the body, two quite distinct localities: (1) Laguna Ubaque,

and principally dorsally situated, there are numer- east slopes of the Cordillera Oriental (2112 m)

The oviducts from Paramo Cruz ous pre-ovarial follicles. arise at Verde (04°34'N 74°02'W)

the wall southeast of Shores of Pedro- outer lateral of each ovary, extending Bogota. (2 ) Laguna

little is east side of Sabana de forwards a way so that there a sperm- palo (2000 m), Bogotâ

O filled tuba anterior the 43'N northwest of to ovary (Fuhrmann, 1914, (04 74°10'W) Bogota.

fig. 6). Dugesia (Girardia) paramensis (Fuhrmann, The penis consists of a stout muscular bulb and 1914) (Figs. 3, 4) a short conical papilla extending into the male

said Fuhrmann Planaria atrium. The vasa deferentia are by paramensis Fuhrmann, 1914: 798.

Euplanaria paramensisr: Kenk, 1930: 292. the where to extend to gonopore, from they recurve Dugesia paramensis: Kenk, 1941: 7. to the bulb. In fact the recurvature can be penis Dugesia ( Girardia ) paramensis: Ball, 1974: 377.

much less (fig. 2). There is no intrabulbar vesicle

Type material. — British Museum (Natural History),

and the narrow duct, which at ejaculatory opens 1928.1.5.87-88, alcohol specimens. BMNH 1928.1.4.258-270,

the tip of the papilla, is more ventrally than slides as follows: 258-260 wholemounts; 261-264 sagittal sec- tions (cf. Fuhrmann, 1914, fig. 9); 265-268 frontal sections centrally located. The male atrium and the penis (cf. Fuhrmann, 1914, fig. 10); 269-270 frontal sections. All

are invested with a nucleate papilla epithelium the available slides have been re-examined but the redescrip-

that the is and based the two series whereas of ejaculatory duct infra- tion new figures are primarily upon

261-264 and 265-268. nucleate.

— The external features The bursa copulatrix is a sacciform organ lying Redescription. anterior to the penis and the bursal canal curves are as described by Fuhrmann (1914: 798) and

the which shows over atrium, bending sharply downwards to specimen 258, a supernumerary eye,

roof wide sinuous duct has redrawn here open into its as a above been as fig. 3. Living specimens

the At the where the bursal canal and dark gonopore. point are up to 20 x 2 mm, jet-black dorsally

the oviducts into The is curves downwards, two open grey ventrally. pharynx pigmented.

it from and these separately behind, beneath are The testes are large, discrete, and numerous

numerous and extensive shell glands (fig. 2). follicles dorsally situated throughout the body-

The bursal canal is lined with a ciliated and length. The first of the testes are immediately

nucleate overlain thin of the be anterior epithelium by layers adjacent (lateral ) to ovaries, or may

circular and The vitellaria subepithelial muscles outer longitu- to them. occur throughout the body

dinal of in the ventral and muscle fibres. The longitudinal muscles but are more extensive half,

The the bursal canal may also extend part way over there are some pre-ovarial follicles. oviducts

of the bursa itself. The vaginal region the bursal show no special peculiarities and enter the bursal canal, between oviducts and atrium, and also the canal separately above the shell glands, in the

wall of the atrium and walls of the usual posterior way.

characterized is in and gonopore, are by a significant The copulatory complex shown fig. 4,

of the inner circular muscle this be with the of Fuhrmann thickening layer. may compared fig. 9

is the The which a drawing of same specimen.

consists muscular and penis of a stout bulb an

Remarks. — In my revision of the Dugesiidae elongate conical papilla. The vasa deferentia

I made of Planaria extend the sides of the and (Ball, 1974) no assignment along pharynx open longistriata. However, it clearly falls within the into the penis bulb with varying degrees of Girardia subgenus as therein defined. Dugesia recurvature (fig. 4; Fuhrmann, 1914, fig. 10),

be all other bifid seminal longistriata may distinguished from to form a vesicle. The wide ejacu- members of the D. duct the of tigrina group (see below) by latory opens almost at tip the penis; its colour pattern, and by its short conical penis Fuhrmann (1914, fig. 9) believed that it opened lacking a developed seminal vesicle. subterminally but he failed to notice the - BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 237

Figs. 1-4. Colombian freshwater triclads of Fuhrmann, 1914, redrawn from the type material. 1, Dugesia longistriata, features BMNH 1928.1.4.245, external drawn from a wholemount; 2, D. longistriata, BMNH 1928.1.4.254,sagittal section of viewed from the left BMNH external features drawn the copulatory apparatus side; 3, D. paramensis, 1928.1.4.258, from a wholemount (same scale as fig. 1); 4,D. paramensis, BMNH 1928.1.4.262, sagittal section of the copulatory

from left apparatus viewed the side (same scale as fig. 2).

= = = = Abbreviations: bc bursa bs bursal ed = male od = copulatrix; canal; ejaculatory duct; go gonopore; ma atrium;

= = shell vd = deferens. oviduct; pe penis; sg glands; vas 238 I. R. BALL - FUHRMANN'S COLOMBIAN PLANARIANS

Fuhr- of in his and I that ture of the tip the papilla specimen (cf. (Borelli, 1898), propose with tall mann's be considered as fig. 4). The male atrium is lined with a species conspecific with flat the latter Kenk, 1974: estai nucleate epithelium, the penis papilla a (cf. 27). Dugesia f

infra- is in the Andes epithelium that is partially or completely a species apparently widespread Ecuador and also into Vene- nucleate; the ejaculatory duct is infranucleate. of Peru, extending has been recorded from The bursa copulatrix lies anterior to the penis zuela and Curaçao. It

Mitchell & Kawakatsu and from it the bursal canal curves smoothly over Colombia by (1973: 658).

the atrium into which it above the opens gonopore.

ciliated infra- cameliae The bursal canal is lined with a Dugesia (Girardia) (Fuhrmann, 1914)

nucleate cuboidal epithelium overlain by thick (Figs. 5-8)

layers of inner circular and outer longitudinal Planaria cameliae Fuhrmann, 1914: 802.

Planaria camaliae Fuhrmann, 1914: 802, lapsus calami. muscle fibres. There is some slight intermingling Euplanaria cameliae: Kenk, 1930: 292. between these two muscle layers. Dugesia cameliae: Kenk, 1941: 7. Dugesia cameria: Mitchell & Kawakatsu, 1973: 658, lapsus

calami. Remarks. In some specimens of this species Dugesia cameriae: Kawakatsu, Hauser & Friedrich, 1976: 215, there are two pharyngés, one in front of the other, lapsus calami.

which Fuhrmann takes to indicate a special form

— British Museum (Natural History), of asexual reproduction. This is discussed fully Type material. 1928.1.5.89-100, alcohol specimens. BMNH 1928.1.4.273-290 in his 1914: (Fuhrmann, 801). Dugesia 277 frontal paper slides as follows: 273-276 wholemounts; sections; the other paramensis may be distinguished from 278 frontal sections; 279-280 sagittal sections (cf. Fuhrmann, 1914: fig. 13); 281 frontal sections; 282 frontal sections; Girardia species with dorsal testes (table I) by its 283-285 sections of 286 frontal sagittal poor quality; sections; conical undivided and large penis papilla, atrium, sections of frontal 287-288 sagittal poor quality; 289-290 sec-

lack of seminal vesicle. tions of All the available slides have been an expanded poor quality. re-

examined but the redescription and new figures are based

primarily on the three series 279-280, 281, and 286.

Distribution. — This species was found

the — in a stream on east slopes of the Cordillera Redescription. The external features are

Paramo Cruz Verde Oriental (3200 m) at as described by Fuhrmann (1914: 803). Living

southeast of to 17 mm and 1 mm wide (04°34'N 74°02'W), Bogota. specimens are up long

have head with and a markedly triangular promi-

nent auricles (fig. 5; Fuhrmann, 1914, fig. 11). Dugesia polyorchis (Fuhrmann, 1914) and The dorsal surface is dark brown or black Planaria polyorchis Fuhrmann, 1914: 802. the ventral surface is As in the other Dugesia polyorchis: Kenk, 1941: 7. pale. species Dugesia (Girardia) polyorchis: Ball, 1974: 376. the pharynx is pigmented.

The testes are numerous follicles situated dor-

Type material. — British Museum (Natural History), and the bodylength, beginning 1928.1.4.271-272, frontal sections on two slides. sally throughout

just behind the ovaries. The vitellaria occur

This in the ventral Remarks. species was discovered by throughout the body, predominantly

Fuhrmann of follicles. The among a sample Dugesia paramensis. half, and there are some pre-ovarial

for Since only one specimen was available study, oviducts show no special peculiarities and enter and since this sectioned before it the bursal canal above the zone was was recog- widely separated

not the in usual nized as new, a satisfactory description was of shell glands, the way.

possible, neither then nor now. Fuhrmann's The copulatory complex is shown in figs. 6-8;

is far it but the 6 of the from which description accurate as as goes fig. being same specimen

sections do show all of the Fuhrmann drew his 13. The consists not copulatory appa- fig. penis

thus the the of muscular rather ratus and affinities of species are a stout bulb and a asymmetric

difficult to assess. It seems to me, however, that papilla projecting into the atrium. In one speci-

everything that can be discerned from the slides men the penis is contracted such that prominent

with is consistent the specimen being folds are 7). The vasa deferentia festaiDugesia present (fig. BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 239

Figs. 5-8. Colombian freshwater triclads of Fuhrmann, 1914, redrawn from the type material. 5, Dugesia cameliae, BMNH

1928.1.4.273, external features drawn from a wholemount; 6,D. cameliae, BMNH 1928.1.4.279-280, sagittal section of the

copulatory apparatus viewed from the right side (same scale as fig. 8); 7, D. cameliae, BMNH 1928.1.4.286, frontal section through the penis viewed from above; 8,D. cameliae, BMNH 1928.1.4.281, frontal section through the copulatory

apparatus viewed from above.

Abbreviations: as in figs 1-4. 240 I. R. BALL - FUHRMANN'S COLOMBIAN PLANARIANS

6 and from Central recurve before entering the penis bulb (figs. & Tahiti (Gourbault, 1978), 24

8 ) and within the bulb they unite, without enlarge- and South America, including some of the Carib-

duct that bean islands. One two have wide distri- ment, to form an ejaculatory opens ven- or species

trally subterminally (fig. 6). The atrium is inter- butions, but most are known from but one or a few

mediate between the totally divided condition of localities.

Dugesia arimana from Trinidad (Ball, 1971, In discussing the South American species, Ev.

and the uniform condition of other Marcus commented the small fig. 3) most (1953) on very

The of between which Dugesia (Girardia ) species. epithelium differences them, makes the un-

the atrium is nucleate and that of the penis and ravelling of their taxonomie relationships difficult.

duct But in the last 25 have ejaculatory is infranucleate. years many more species

been and in variation The bursa copulatrix lies anterior to the penis described some patterns and it from the bursal canal curves smoothly over begin to appear.

the atrium towards the The The difficulties at the level notwith- gonopore. epithelium species of the bursal canal is infranucleate and is overlain standing, it is possible to divide the species on the

thick of circular clear by a musculature inner and outer basis of two characters, the distribution of

There is the and the longitudinal fibres. some intermingling testes course of the bursal canal. It

of In of that the charac- of the layers musculature. many the slides must be admitted, however, two

ters and spermatophores were present in the bursa copu- are not completely congruent the respec-

latrix. tive made groups are not quite disjoint.

The question as to the reliability of the position

— In revision of the character has Remarks. my Dugesiidae of the testes as a taxonomie often

(Ball, 1974) I made no assignment of Planaria been debated (De Beauchamp, 1939; Ball, 1971;

cameliae. falls within the of the However, it clearly sub- Kenk, 1975). Dugesiidae Dugesia gono-

Girardia therein defined. Of the Girardia the genus as cephala group, subgenus Dugesia Ball, 1974,

the species with dorsal testes (table I) Dugesia came- distributed throughout Old World, are charac-

liae is most similar to D. miltgeni in its markedly terized by their numerous dorsal testes. Until

ventral subterminal opening of the ejaculatory recently most Girardia species differed in their

and in the of the numerous situated duct, histological structure copu- closely packed ventrally testes,

latory apparatus. The penis of the Colombian and dorsal follicles were the exception rather than

is and the the With it has species more pointed, however, vasa rule. more recent work, however,

deferentia recurve before entering it. now become clear that no less than 14 out of the

30 species have dorsal testes (table I). Moreover,

— Fuhrmann it is of Distribution. According to noteworthy that all but one these species

rivers close I called (1914: 802) this species was found in have what have a smoothly curved bursal

to the city of Medellin (06°15'N 75°36'W) in canal.

the Cordillera Central (1540-1820 m). Guaca The type species of the subgenus, Dugesia

presumably refers to the Rio Guaca (06°15'N (Girardia) tigrina, and the species most widely

I have canal 75°52'W); been unable to trace Cafetal distributed, possesses a bursal that curves

the La Camelia. over copulatory apparatus and then bends

sharply downwards at the point of entry of the SYSTEMATIC RELATIONSHIPS OF THE oviducts (Ball, 1971, figs. 5 & 7). This character COLOMBIAN SPECIES is and constant in related present many species

Freshwater planarians of the family Dugesiidae such as Dugesia longistriata (fig. 2). The alter-

the occur throughout New World (Ball, 1974, native condition is a smooth, uninterrupted, down-

1975). Most of the New World forms belong to ward curvature of the bursal canal (figs. 4 & 6).

the With there correlation genus Dugesia, subgenus Girardia, and at one exception is a between

there 30 of and present are some valid species known; the presence dorsal testes the smooth con- five from continental North America, one from ditionof the bursal canal (table I). It is also note- bijdraGen tot De dierkunde, 50 (l) - 1980 241

Table I

List of Girardia species with dorsal testes.

Species Atrium 1 Bursal canal 1 Distribution Source of data

1 andina uniform smooth Andes, Argentina Type material

2 chilla (un) divided 2 angled/smooth 2 Central & Southern Marcus, 1954;Hyman, 1959

Chile

3 3 arizonensis divided smooth Arizona, USA Kenk, 1975

4 arimana divided smooth Trinidad/S. Vincent Ball, 1971; Kawakatsu et al., 1976

Venezuela

divided smooth in 5 miltgeni Guadeloupe Goui'balt, press

6 antillana uniform smooth Puerto Rico Kenk, 1941

7 cubana uniform smooth Cuba Codreanu & Balcesco, 1973

8 barbarae uniform smooth Mexico Mitchell & Kawakatsu, 1972

9 typhlomexicana uniform smooth Mexico Mitchell &: Kawakatsu, 1972

10 mckenzei uniform smooth Mexico Mitchell & Kawakatsu, 1973

± uniform 11 cam el iae smooth Colombia Type material

12 paramensis uniform smooth Colombia Type material

13 veneranda uniform smooth Brasil Martins, 1970

14 hypoglauca divided angled Brasil Marcus, 1946

1 of these characters the For an explanation see main text.

2 alternatives Marcus (1954) shows the bursal canal as angled and the atrium divided; Hyman (1959) gives the

3 The bursal canal is sinuous and approaches the angled condition (Kenk, 1975, fig. 5).

D. D. D. worthy that no less than 10 of the species listed arizonensis, arimana, and miltgeni have

in table I (3-12) originate from a geographical a clearly divided atrium that is unusual in the

The of all area embracing Mexico and contiguous U.S.A., Girardia species. evolutionary history

Central the and northern these forms be America, Caribbean, may be expected to clarified by the

South America. material collected on the University of Amsterdam

Of the Colombian species redescribed here, Expeditions to the West Indian Islands by Pro-

Dugesia longistriata and D. festai ( = polyorchis) fessor Dr. J. H. Stock and Dr. L. Botosaneanu, belong clearly with the tigrina-group. The two and which will form the subject matter of a future other The of species, D. cameliae and D. paramensis, are report. relationships certain more southern members of that have D. D. D. and a large subgroup appears to forms, andina, chilla, veneranda, its centre of diversification in the Central Ameri- D. to the Caribbean hypoglauca (table I) group

and Caribbean This shows must await further assessment can area. group even (cf. Ball, 1971; further diversification in that certain of the species, Kawakatsu et al., 1976).

ACKNOWLEDGEMENTS

I I am grateful to both Mr. S. Prudhoe and Dr. D. I. Gibson visit to the Museum was made while was the recipient of for their visit the British Museum Grant A0016 of the National Research hospitality during my to Operating Council of (Natural History), and to them and the Trustees of the Canada. I am also grateful to my wife, Maria Tran Thi

BMNH for lending me the specimens for further study. My Vinh-Hao, for preparing the figures. 242 I. R. BALL - FUHRMANN'S COLOMBIAN PLANARIANS

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Received: 13 March 1980