identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E95D02FFF3FFAEC8F4FF021A90FEEC.text	03E95D02FFF3FFAEC8F4FF021A90FEEC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phagocata punctata Bromley-Schnur 2021	<div><p>Phagocata punctata sp. nov.</p> <p>(Figs 1, 2A, 3 – 5; Tables 1 – 5)</p> <p>Material examined. Holotype. HUJ.Kin.1, sagittal sections on 3 slides; Paratypes: HUJ.Kin.2, sagittal sections on 3 slides; HUJ.Kin.3, transverse sections on 7 slides. From laboratory culture, collected in the field, June 1972. Material deposited in the Zoological Collections of the Department of Ecology, Systematics and Evolution, Hebrew University, Jerusalem, Israel.</p> <p>Type locality. <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=35.638203&amp;materialsCitation.latitude=32.764687" title="Search Plazi for locations around (long 35.638203/lat 32.764687)">Two</a> adjacent sites in the littoral water off the south-eastern shore of Lake Kinneret: N32.764688 E35.638202.</p> <p>Habitat. The type locality of Lake Kinneret where Ph. punctata was found lies in the River Jordan Rift Valley and is presently about 209 m below sea level. Ph. punctata was usually found in clusters in small crevices on the undersides of small pitted rocks and boulders at a depth of 50 to 150 cm. These rocks typically also bore patches of a sponge, Cortispongilla barroisi (Topsent, 1892) and a bryozoan, Fredericella sultana Blumenbach, 1779. No other triclads were found at these littoral sites.</p> <p>The lake water is subject to considerable seasonal temperature fluctuations of between 14 and 29 oC and the average salinity measured in 1972–3 was around 230–240 mg /l Cl´. However, in the past, both the water level and the salinity have fluctuated greatly, the water level ranging from a maximum of about 209 m to about 215 m below sea level (Markel 2014) and the salinity ranging from about 200 mg /l Cl´ to a peak of 390 mg /l Cl´ in the early 1960s. In the mid-1960s, considerable amounts of saline water were diverted into a salt-water channel from western saline springs at the edge of the lake to the lower Jordan River, and subsequently, the salinity levels of the lake decreased by about 30% (Rimmer &amp; Nishri 2014). In 2018, after several years of drought as well as increased irrigation diversion for agriculture, historically low levels of water in the lake, to - 213 m, resulted, and the salinity again rose to almost 300 mg /l Cl´, one of its highest levels in 50 years. By spring 2020, after an exceptionally rainy winter, the lake level again rose by just over 3 m to almost - 209 m.</p> <p>Diagnosis. Phagocata punctata sp. nov. is characterized by grey or grey-brown spots on a light grey background dorsally, very light or white ventrally. It has a large distinct penis bulb with a short blunt, conical papilla. There is a blind-ending caecum in the penis, lying ventrally to the ejaculatory duct, extending back into the penis bulb where it has an enlarged lumen. The caecum opens into the distal part of the ejaculatory duct. Chromosome portrait: diploid (2n = 34), with the first chromosome pair about twice as long as the second pair.</p> <p>Etymology. The specific epithet refers to the spotted or speckled appearance of the new species.</p> <p>Description. The main external features of Ph. punctata are shown in Table 1, where they are compared with those of Ph. armeniaca, a description of which follows that of Ph. punctata.</p> <p>There are numerous small pigmentation granules beneath the basement membrane of the dorsal epithelium and large intermittent clusters of pigment underlying the dorsal musculature. The pharynx is composed of two muscle layers in the outer muscle sheet. Both the outer, longitudinal layer and the inner circular layer are about 2.5 µm thick. Cyanophilous glands are seen in the head region under the dorsal musculature, extending posteriorly about halfway to the pharynx insertion and anteriorly to the tip of the head. They are well developed ventrally in the median body region and also around the ovaries where they extend to the dorsal body wall. There are also many cyanophilous glands in the region of the pharynx insertion. The mouth opens just beyond the posterior end of the pharyngeal pouch.</p> <p>The ovaries are situated ventrally, just behind the brain, at about 1/6 th of the distance between the brain and the root of pharynx, after the first main lateral digestive branch. They are round to oval in shape, measuring 100–175 µm dorso-ventrally and 100–120 µm antero-posteriorly. The oviducts leave the ovaries from a mid-lateral position and lie above the ventral nerve cords. The right oviduct passes on the right side of the penis while the left one lies between the penis and the bursal canal. The common oviduct enters the bursal canal dorsally, near its junction with the male atrium.</p> <p>The testes have a ventral position, but may extend into the dorsal part of the body as well. The testes extend from the posterior level of the ovaries almost to the tip of the tail. They are round to pear-shaped, 150–200 µm in diameter, lying in two rows, one on either side of the body.</p> <p>The copulatory bursa is large, sacciform and thin-walled, measuring 200–300 µm dorso-ventrally and 200–400 µm antero-posteriorly. It often contains one or more spermatophores. The spermatophore bulbs measured around 150 µm and the stalks up to 400 µm. The bursal canal exits from the dorsal posterior part of the bursa and lies dorsally, to the left of the penis. The muscle sheath of the canal is of uniform thickness and consists of a thin layer of circular muscle surrounded by an outer longitudinal muscle layer. It is lined with an epithelium of tall, columnar ciliated cells with basal nuclei. The bursal canal dilates near the atrium. The female, male and common atria are distinct. The male atrium has a dorsal epithelium of cuboid cells and a ventral epithelium of flattened cells. It is surrounded by a layer of subepithelial circular and then longitudinal muscle fibres. The female atrium lies to the left of the common atrium and as a diverticulum of it. A group of eosinophilic cells, presumed to be shell glands, enter the common atrium. There is a short gonoduct, about 100 µm in length (Figs 3, 4).</p> <p>There is a large distinct penis bulb with a small and blunt, conical papilla. There is a ring of outer longitudinal and inner circular muscle near the papilla tip. The vasa deferentia enter the penis antero-laterally and open into a distinct round or oval seminal vesicle lined with smooth flat epithelium and surrounded by circular muscle. There is a muscular constriction in the globular seminal vesicle which partially divides the seminal vesicle into two parts. The ejaculatory duct exits from the distal part, forming a narrow tube opening terminally (Figs 3, 4).</p> <p>The most outstanding characteristic of the reproductive system of Ph. punctata is the caecum, a blind-ending, kidney-shaped tube lying ventrally to the ejaculatory duct in the penis papilla and opening into the distal portion of the ejaculatory duct. The caecum extends back the whole length of the penis papilla and into the penis bulb where it has an enlarged lumen at its blind end. The lumen of the caecum and of the ejaculatory duct become continuous at the tip of the penis papilla.</p> <p>Karyology of Ph. punctata. Thirty mitotic metaphase plates from five specimens studied revealed 17 pairs of chromosomes. Figure 5 shows three sets of chromosomes, two of which show the presence of a small supernumerary chromosome, about ½ the size of chromosome 17. The first pair of chromosomes are significantly larger than all the others, with number 2 being approximately half the size of number 1. Number 3 is noticeably smaller than number 2 and pairs 4 to 17 decrease gradually in size, although several pairs are indistinguishable. Pair 17 is about 1/5 th the size of pair one.</p> <p>Chromosome 1 is large and metacentric with the highest centromeric index value. Chromosomes 2, 3, 4, 7, 10 &amp; 11 are all submetacentric, but number 3 is borderline submetacentric-subtelocentric. All other chromosomes are metacentric or borderline metacentric-submetacentric. The standard deviations for the centromeric indices are rather high in many cases.</p> <p>Tables 2 and 3 show a comparison of mean values and standard deviations of the relative lengths and centromeric indices of both species of Phagocata.</p> </div>	http://treatment.plazi.org/id/03E95D02FFF3FFAEC8F4FF021A90FEEC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Bromley-Schnur, Heather J.	Bromley-Schnur, Heather J. (2021): The genus Phagocata Leidy (Platyhelminthes, Tricladida) in Israel, a new species of Phagocata from Lake Kinneret, and an emended description of Dugesia salina. Zootaxa 4969 (2): 293-317, DOI: https://doi.org/10.11646/zootaxa.4969.2.4
03E95D02FFF7FFA7C8F4FA791A01FA84.text	03E95D02FFF7FFA7C8F4FA791A01FA84.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phagocata armeniaca (Komarek 1916)	<div><p>Phagocata armeniaca (Komárek, 1916)</p> <p>(Figs 1, 2B, 6 – 8; Tables 1 – 5)</p> <p>Material examined. Material was collected from the spring sites of Dan, N 33.248824 E35.652379 and Banias, N33.248663048 E 35.695301297 in northern Israel. Slide preparations examined were from Banias as follows: HUJ. Ban.3, sagittal sections on 4 slides; HUJ.Ban.4, transverse sections on 7 slides; HUJ.Ban.5, transverse sections on 15 slides; HUJ.Ban.6, sagittal sections on 5 slides. From laboratory culture, collected in the field, June 1972. Material deposited in the Zoological Collections of the Department of Ecology, Systematics and Evolution, Hebrew University, Jerusalem, Israel.</p> <p>Habitat. The spring habitats of both Dan and Banias in northern Israel, where Ph. armeniaca was found, have water of an almost constant temperature of around 15–16 oC and the average salinity was 8–12 mg /l Cl´ at Dan and 12–16 mg /l Cl´ at Banias. The water is mainly fast flowing over a stony substrate, but some stones were partly submerged in sand or silt. The water depth where the triclads were found was usually in the range of 10 to 30 cm. Dan is a large complex of springs and streams and there are areas of slower flowing water adjacent to the main river. Other Tricladida found at these sites were Dendrocoelum dani Bromley, 1982, Dugesia golanica Bromley &amp; Benazzi, 1991 and Dugesia biblica Benazzi &amp; Banchetti, 1973, now considered to be a junior synonym of Dugesia sicula Lepori, 1948 (Solà et al. 2015).</p> <p>Description. Comprehensive descriptions of Ph. armeniaca have been made by both Komárek (1916) and by De Beauchamp (1958). The description following here is intended to point out several differences in the reproductive apparatus between the two Phagocata species found in northern Israel. Table 1 shows a comparison of the external morphology of the two species. Many of the internal features are shared by both species, and in order to avoid unnecessary repetition, the description will be compared with that of Ph. punctata in some cases.</p> <p>The pigmentation of Ph. armeniaca is seen dorsally as numerous granules scattered throughout the mesenchymal cells underlying the dorsal muscle layer, and fewer granules in the ventral mesenchyme.</p> <p>The ovaries are situated ventrally, just behind the brain, at about 1/8 th of the distance between the brain and the root of the pharynx. The pharynx is composed of two muscle layers in the outer muscle sheet. The outer, longitudinal layer is about 2.5 µm thick and the inner, circular layer is about 7.5 µm thick. The mouth is located just posterior to the hind end of the pharyngeal pouch.</p> <p>Cyanophilous glands are abundant, especially in the anterior quarter of the body, dorsally and ventrally, between the gut branches and the mesenchymal cells internal to the body wall musculature. They are also very prominent anterior to the eyes, in a median region between the eyes and the brain, as well as in the central region between and behind the ovaries, extending back as far as the first and second testes. They were also seen in the central body region immediately anterior to the pharynx insertion.</p> <p>The female reproductive system is essentially similar to that described above for Ph. punctata.</p> <p>The position and extent of the testes are similar in both species of Phagocata. However, there are significant differences in the details of the male reproductive systems between the two species.</p> <p>Ph. armeniaca has a small penis bulb, while the papilla is large, cylindrical and pointed, with a ring of circular muscle almost at the tip. The vasa deferentia enter the lateral walls of the penis bulb. The seminal vesicle is represented by a wide tube, lined by very tall cells, which almost fill the lumen of the penis bulb and anterior part of the papilla where the two vasa deferentia enter. The seminal vesicle is constricted about 2/3 along its length by an eosinophilous plug of cells which apparently separate it from the ejaculatory duct, which opens terminally (Figs 6, 7).</p> <p>Similar to that described above for Ph. punctata, the penis of Ph. armeniaca has a blind-ending, kidney-shaped caecum lying ventrally to the ejaculatory duct in the penis papilla. The caecum extends back the whole length of the penis papilla and slightly into the penis bulb, remaining narrow for its whole length. The lumen of the caecum and of the ejaculatory duct become continuous at the tip of the penis papilla in both species.</p> <p>Karyology of Ph. armeniaca. Mitotic metaphase plates were studied in five specimens, and 17 pairs of chromosomes were clearly counted in 25 metaphase plates. Figure 8 shows three sets of chromosomes. The first pair of chromosomes is significantly larger than all the others, and is about twice the length of pair 2. Number 3 is noticeably smaller than number 2 and pairs 4 to 17 decrease gradually in size, although several pairs are indistinguishable. Pair 17 is about 1/5 th the size of pair one.</p> <p>Chromosome 1 is large and metacentric with the highest centromeric index value. Chromosomes 2, 3, 4, 7 and 11 are all submetacentric. All other chromosomes are metacentric or borderline metacentric-submetacentric. The standard deviations for the centromeric indices are rather high in many cases.</p> <p>Tables 2 and 3 show a comparison of mean values and standard deviations of the relative lengths and of centromeric indices of both species of Phagocata.</p> <p>Reproductive behaviour of the two Phagocata species. Collections of Ph. armeniaca at all seasons from both</p> <p>Dan and Banias included young, immature as well as larger, sexually mature animals. At Banias, where the population was larger and easier to monitor, cocoons were found on stones bearing mature animals during both summer and winter. The cocoons were round to oval in shape, unstalked and attached to the substrate such that the long axis of an oval cocoon was always parallel to the substrate. No evidence of fission was ever seen and presumably sexual reproduction occurs throughout the year.</p> <p>Ph. punctata was first discovered in Lake Kinneret during the spring and summer of 1972 (April to September) when only asexual animals were found with no evidence of fission having occurred. The specimens were up to 7 mm in length, a size normally associated with mature specimens of Ph. armeniaca. However, during the winter months (November to March), both immature and sexually mature animals were found, and on a few occasions, cocoons were found on stones also bearing mature planarians. It thus appears that conditions suitable for sexual reproduction of Ph. punctata occur only when the water temperature is 20 oC or lower.</p> <p>In the laboratory, both species of Phagocata were very easy to maintain and breeding was established at 18 oC. Asexual reproduction was never observed although the regenerative powers of both species, when cut, was good. Comparative data on cocoons and emergence of young at 18 oC is given in Table 4.</p> <p>The cumulative rate of cocoon production of 10 animals of each species, kept separately, was followed for 100 days at temperatures of 15 oC, 18 oC and 20 oC rising gradually to 25 oC. In both species, cocoon production was greatest at 18 oC, lowest at 15 oC (the temperature at Banias) and an intermediate rate at 21–23 oC, which was the first phase of the high temperature period (Table 5). Cocoon production slowed down greatly or stopped and cocoons already deposited did not hatch at temperatures above 23 oC (Ph. armeniaca) and 25 oC (Ph. punctata) at which temperature some animals of both species began to break up. By day 100, the cumulative egg production of the Ph. punctata animals was approximately twice that of the Ph. armeniaca animals, and that of the F 1 cross was intermediate (Table 5).</p> <p>The following observation indicates that the induction of sexual maturity in asexual animals of Ph. punctata is temperature dependant. Asexual animals were collected from the field in May, 1973, when the water temperature was 25 oC. Half the animals were cultured at 18 oC and the others at room temperature of 20–28 oC. After 3 months, the 18 oC animals were all sexually mature and depositing cocoons, while none of the room temperature animals had matured. When the latter were transferred to 18 oC, they matured within a few weeks.</p> <p>Cross-breeding experiments. Since the morphology and karyology of both species of Phagocata were very similar, an attempt was made to cross-breed them. Immature animals of each species were raised in isolation to maturity at 18 oC and then one specimen of each, per jar, was cultured together. Both parental types deposited fertile cocoons. The cocoons were often larger than either parental type (ranging from 1.2 x 0.9 to 1.5 x 0.9 mm), and the number of young emerging was generally higher (5–17 per cocoon, usually 7–10). The hatching time was approximately the same (16–21 days) as was the range in size of emerging young (0.5–2.75 mm).</p> <p>Most of the F 1 animals were normal and healthy and the examination of 100 animals showed a range of gut branching patterns, i.e., fairly branched like Ph. armeniaca, fairly sparse like Ph. punctata, or intermediate (in a ratio of 2:2:1). Likewise, after a few days when pigmentation patterns became clear, the range of patterns of the F 1 animals was noted, with some unspotted, some definitely spotted and the majority very slightly spotted, (in a ratio of about 1:1:3). However, later examination showed that very few animals acquired the full spotted pigmentation patterns typical of adult Ph. punctata, thus the ratio changed to 4:1:7 (unspotted like Ph. armeniaca, definitely spotted like Ph. punctata and intermediate).A small percentage of the F 1 animals (about 0.5%)were joined at the head or anterior body region like “Siamese twins”. Such abnormalities were never observed among the pure-bred offspring of either parental type.</p> <p>Some of the F 1 animals, with a random assortment of pigmentation patterns were left to grow to maturity, and at 7–8 weeks old, the first cocoons were deposited. Cumulative cocoon production of 10 mature F 1 animals was also determined at various temperatures over a period of 100 days as with the pure-bred animals described above. The results are shown in Table 5, and indicate a cocoon production at all temperatures intermediate between those of the two parent types. The F 2 animals showed a range of pigmentation from brown, to brown speckled, pale grey and clearly grey spotted. From the 117 cocoons that hatched during the experimental period, 1065 young were produced, an average of 9.1 per cocoon. Most were normal, but there were 5 “Siamese twins” amongst them.</p> <p>A separate set of F 1 animals, with a range of pigmentation patterns, was kept for observation and breeding at 18 oC. All animals were mature and measured 8–11 mm in length. 219 cocoons were deposited producing 1279 young (average 5.8 young per cocoon). Again, most of the F 2 offspring (89%) were normal, while the remaining 11% included 15 “twins” representing 1% of the total, as well as a new type of abnormality, small round or oval, pale brown “monsters”, 0.5–1.5 mm in length. There were 128 of these monsters (10% of the total F 2 offspring); they were mainly 2-eyed, with no pigment-free area around the eyes; one had a single eye, one had five eyes; they apparently had a degenerate pharynx and could not eat or grow, and after 15 to 30 days of gliding around the culture jars, they died.</p> <p>Discussion. While the external features and the habitats of the two species of Phagocata from Israel are very different and completely isolated from each other in nature, it is clear from the above morphological considerations that there is a very close relationship between Ph. armeniaca and Ph. punctata. However, although they share the presence of a penial caecum, they can easily be distinguished by the significant differences in the shape of the both the penis bulb and the penis papilla: large bulb and a short conical papilla in Ph. punctata and small bulb and a large cylindrical papilla in Ph. armeniaca.</p> <p>Tables 2 and 3 show the close karyological similarity between the two species. The relative lengths of the chromosomes in the two species are very similar. The main differences are in the centromeric indices of chromosomes 3 and 10. Chromosome 3 in Ph. armeniaca is clearly submetacentric while in Ph. punctata it is submetacentric bordering on subtelocentric; chromosome 10 in Ph. armeniaca is metacentric, while in Ph. punctata it is submetacentric. In addition, two of the metaphase plates examined in detail for Ph. punctata showed the presence of a small supernumerary chromosome, but there is not enough data to know if this is a constant feature.</p> <p>The taxonomic position of Ph. armeniaca with regard to other European representatives of the genus was discussed in detail by De Beauchamp (1958). It is noteworthy that De Beauchamp pointed out that in the original description of the penis of Ph. armeniaca by Komárek (1916), he mistakenly believed that the caecum was in fact one of the vasa deferentia, and that both ran independently through the length of the penis. He also notes that Komárek’s material was collected from stenothermic mountainous springs and streams in Armenia, perhaps not too dissimilar to the cold streams of Dan and Banias in Israel. In contrast, De Beauchamp’s material was collected from lakes in eastern Turkey and presented a quite different type of habitat. He suggested that the planarians living in the lakes may become sexually mature either in colder springs nearby or only in the winter. This is the situation with Ph. punctata, which becomes mature in Lake Kinneret only at winter temperatures lower than about 20 oC.</p> <p>The systematics of the genus with reference to five new pigmented species from Spain have been discussed more recently by Sluys et al. (1995) and Vila-Farré et al. (2011). Most animals now assigned to the genus Phagocata are small white animals with a truncated head and include the former junior synonyms of Albiplanaria Komárek, 1926, Fonticola Komárek, 1926, and Penecurva Livanov &amp; Zabusova, 1940 (Sluys et al. 1995). However, apart from Ph. armeniaca, there are several other pigmented species of Phagocata in Europe, including the five from Spain (Ph. ullala Sluys, 1995, Ph. flamenca Vila-Farré &amp; Sluys, 2011, Ph. asymmetrica Vila-Farré &amp; Sluys, 2011, Ph. gallaeciae Vila-Farré &amp; Sluys, 2011, Ph. pyrenaica Vila-Farré &amp; Sluys, 2011) as well as the endemic species of Lake Ohrid, Ph. maculata (Stanković, 1938). These six pigmented species are all quite distinct from Ph. armeniaca and Ph. punctata regarding the penis papilla, which in both these latter species have a blind-ending caecum opening into the distal part of the ejaculatory duct. The only other Phagocata species known to have such a diverticulum is the pigmented North American Ph. velata (Stringer, 1909), which differs from the Israeli species in other morphological features and especially in its karyological pattern of 60 to 96 chromosomes, corresponding to different degrees of polyploidy (Whitehead 1965).</p> <p>The number of chromosomes for several species of Phagocata have been described (see Sluys et al. 1995). Among these, Ph. vitta (Dugès, 1830) and Ph. paravitta (Reisinger, 1923), both unpigmented species, may have a diploid complement of 34, like Ph. armeniaca and Ph. punctata, although Ph. vitta may also be polyploid. The pigmented species, Ph. ullala and Ph. pyrenaica also have a diploid complement of 34 chromosomes, and like Ph. vitta, as well as Ph. armeniaca and Ph. punctata, the first pair of chromosomes are about twice the length of the second pair, acting as marker chromosomes (Sluys et al. 1995). However, both Ph. ullala and Ph. pyrenaica have chromosome pairs 2 and 3 of about the same length. In contrast, with both Ph. armeniaca and Ph. punctata, chromosome pair 2 is clearly longer than pair 3 (see Table 2).</p> <p>The differences in reproductive behaviour of the two Phagocata species is probably a reflection of the different field conditions. At each temperature, the cocoon production of Ph. punctata was approximately twice as high as that of Ph. armeniaca, which may indicate a higher fecundity of the animals in Lake Kinneret under field conditions, possibly as an adaptation to the shorter breeding period in nature during the winter months.</p> <p>The cross-breeding experiments showed that despite the morphological differences of the two types of Phagocata, they can, under laboratory conditions, produce mostly fertile offspring. However, the relatively high proportion (up to 11%) of abnormalities amongst the F 2 animals indicates an incomplete reproductive compatibility, favouring the separation at species level. Dahm (1958, page 117) has noted that “The abundance of evidently near-related but still isolated forms (of triclads) is a feature characterizing those animals which are said to represent “young species”, still being in a rapid stage of evolution”. It seems probable that this is the situation with the separation and speciation of the Lake Kinneret animals from the more widely distributed Ph. armeniaca stock and which has taken place in relatively recent geological times.</p> </div>	http://treatment.plazi.org/id/03E95D02FFF7FFA7C8F4FA791A01FA84	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Bromley-Schnur, Heather J.	Bromley-Schnur, Heather J. (2021): The genus Phagocata Leidy (Platyhelminthes, Tricladida) in Israel, a new species of Phagocata from Lake Kinneret, and an emended description of Dugesia salina. Zootaxa 4969 (2): 293-317, DOI: https://doi.org/10.11646/zootaxa.4969.2.4
03E95D02FFFEFFBFC8F4F97F1D17FEEC.text	03E95D02FFFEFFBFC8F4F97F1D17FEEC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dugesia Girard 1850	<div><p>Genus Dugesia Girard, 1850</p> <p>Material examined. Neotype: HUJ.Oct.1, Sagittal sections on 18 slides.</p> <p>Other material: Transverse sections on 24 slides (HUJ.Oct.2), sagittal sections on 11 slides (HUJ.Oct.3) and transverse sections on 11 slides (HUJ.Oct.5). Material was collected during the summers and winters of 1971–1974 and ex-fissiparous specimens were obtained in the laboratory. The material is deposited in the Zoological Collections of the Department of Ecology, Systematics and Evolution, Hebrew University, Jerusalem, Israel.</p> <p>Type locality: En Sheva (Tabgha), N32.873166 E35.548973, a saline spring on the north-western shore of <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=35.548973&amp;materialsCitation.latitude=32.873165" title="Search Plazi for locations around (long 35.548973/lat 32.873165)">Lake Kinneret</a>, also known as Lake Tiberias.</p> <p>Diagnosis. Dugesia salina is characterized by occupying a very distinct saline habitat. It has a low triangularshaped head with hardly discernible auricles; it is pale brown or grey in colour dorsally, and very pale or white ventrally. Ex-fissiparous specimens have a large well-developed penis, which is situated to the left and slightly anterior to the larger adenodactyl which has a well-developed lumen in the distal part. The penis papilla is large and cylindrical and blunt at its tip where it bends ventrally. There is a distinct funnel-shaped diaphragm separating the seminal vesicle and the ejaculatory duct; the opening of the ejaculatory duct is terminal. Chromosome portrait: diploid, 2n = 16.</p> <p>Habitat notes. En Sheva is a complex of several springs of varying salinities, and Dugesia salina was found on the undersides of basalt rocks in a small spring-fed pool about 5 m south of an octagonal building enclosing the largest spring. The salinity was in the region of 1500 mg /l Cl´ and the temperature was 25–27 oC.</p> <p>Dugesia salina was also found at two other saline locations, namely, En Soda, N32.518836777 E35.534845127, a saline spring several km south of <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=35.57061&amp;materialsCitation.latitude=32.67744" title="Search Plazi for locations around (long 35.57061/lat 32.67744)">Lake Kinneret</a> (salinity ± 1200 mg /l Cl´, temperature 24–27 oC), and in the River Jordan near its outlet from <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=35.57061&amp;materialsCitation.latitude=32.67744" title="Search Plazi for locations around (long 35.57061/lat 32.67744)">Lake Kinneret</a>, N32.677439367 E35.570610941 (temperature 15–26 oC). <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=35.57061&amp;materialsCitation.latitude=32.67744" title="Search Plazi for locations around (long 35.57061/lat 32.67744)">This</a> latter site is where water from several saline springs on the western lake shore (including En Sheva), which prior to 1965 drained directly into the Lake, are now collected in a saltwater channel (the saltwater carrier) and discharged into the River Jordan south of the Lake. No other triclads were found at any of these three sites.</p> <p>Animals were collected from the field during all seasons of the year from En Sheva and En Soda and all were asexual, measured 1.5–8.5 mm, and most showed evidence of recent fission. Those collected from the River Jordan during winter months at 15 oC were slightly larger (6–8.5 mm) and did not show any signs of recent fission.</p> <p>The animals from En Sheva and En Soda were difficult to maintain in the laboratory at 18 oC or 23 oC and many died within a few days of collection, while those from the River Jordan reproduced freely by fission at 18 oC, but none were ever observed to mature. However, several specimens from the type locality of En Sheva, which were kept at a fluctuating room temperature of 12–25 oC, did mature over a period of 3–9 months after collection. A constant temperature of 18 oC or 23 oC did not induce maturation. Saline water from the original habitats was not a prerequisite for laboratory survival or maturation as the animals could be kept satisfactorily in tap water.</p> <p>Description. The planarians from all three populations were similar in external appearance, the only difference being in a greater degree of speckling in the dorsal pigmentation among the animals from the River Jordan. The following description is based mainly on ex-fissiparous animals from En Sheva. The animals have a low triangularshaped head with hardly discernible auricles, which merge gradually into the body region. They are pale brown or grey dorsally and very pale or white ventrally. The animals from River Jordan have a pale or mid-brown mottled colour dorsally and are pale ventrally. After laboratory culture, some animals attained a length of up to 15 mm.</p> <p>The dorsal epithelium is about 15 µm thick and contains many rhabdites, while the ventral one is about 7.5 µm thick with very few rhabdites. The outer muscle sheath of the pharynx consists of two layers only, an outer, subepithelial longitudinal layer 4–5 µm thick, followed by a circular layer, 5–7.5 µm thick. Minute pigment granules are scattered amongst the dorsal body wall muscle layer and its underlying mesenchyme. The inner epithelium of the pharynx is underlain with a thick layer of circular muscle followed by 2–3 layers of longitudinal muscle.</p> <p>The eyes are placed just anterior to the auricles and are surrounded by a fairly large unpigmented area. They are slightly closer together than their distance from the lateral margins of the head and from the anterior tip of the head. The eyes may attain a size of 180 µm in antero-posterior direction. The intestine is highly branched in large specimens. In smaller animals, there are 5–6 lateral branches on the anterior ramus and about 9 lateral branches on the posterior rami. The anterior intestinal trunk forms an unbranched extension in the midline, reaching between the eyes and sometimes just anterior to the eyes (Fig. 9).</p> <p>The insertion of the pharynx in an asexual animal just prior to division is almost halfway down the body and its length is ¼ to ⅕ of the total body length. In laboratory cultured ex-fissiparous animals, the pharynx occupies a more forward position and its length is about 1/7 th of the total body length. The mouth opens slightly posterior to the end of the pharyngeal cavity. The mouth is just over halfway down the body and the gonopore occupies a position just over ¾ down the body length.</p> <p>The two ovaries lie ventrally, just internal to the ventral nerve cords, between the 2 nd and 3 rd or 3 rd and 4 th lateral intestinal branches. They are located just behind the brain, at about one-fifth to one-sixth of the distance between the brain and the root of the pharynx. The ovaries are round to oval in shape and may attain a maximum diameter of 140 µm. The oviducts lead out laterally from the posterior end of the ovaries and open separately, at more or less the same level, into the bursal canal at its junction with the common atrium. The left oviduct enters the base of the bursal canal ventro-laterally, while the right oviduct enters dorso-laterally.</p> <p>The bursa is large and laterally compressed and takes up much of the dorso-ventral space of the body. It attains a lateral diameter of 150–200 µm and a maximum dorso-ventral diameter of about 300 µm. The bursal canal emerges from the posterior part of the bursa and lies dorsally, to the left of the midline, overlying the penis. The glandular, nucleated epithelium of the bursal canal is surrounded by a thin layer of longitudinal muscle and an outer thicker layer of circular muscle. A few extra longitudinal fibres of ectal reinforcement can be seen where the bursal canal enters the common atrium. The canal remains narrow here with hardly any dilatation. The epithelium is ciliated with basal nuclei. Shell glands open into the bursal canal from both anterior and posterior directions in the region of the oviduct openings.</p> <p>The testes are principally dorsal and begin at the level of the ovaries. They continue posteriorly to within 550 µm from the posterior tip of the tail. Pre-pharyngeally, there are 3–4 testes across each longitudinal row; in the pharyngeal and copulatory regions there are 1–3 across each row, and posterior to the gonopore, there are single testes across each row. The testes are oval, flattened laterally and reach up to 130 µm in maximum diameter. Spermatogenesis in these laboratory matured animals appeared normal, with the development of normal sperm. The vitelline glands are predominantly ventral in the mesenchyme, but some are also situated dorsally, between the testes.</p> <p>The copulatory apparatus consists of a large and well-developed penis, which is situated to the left and slightly dorsal and anterior to the larger adenodactyl. The penis bulb is 200–250 µm in diameter and is composed of alternating layers of longitudinal and circular muscle externally. The central tissue, anterior to the seminal vesicle, consists of parenchyma traversed by diagonal muscle fibres. The vasa deferentia expand to form spermiducal vesicles about halfway down the pharynx length. After having decreased in diameter, the vasa deferentia enter the lateral walls of the penis bulb, more or less symmetrically, just anterior to where the penis bulb and the adenodactyl bulb appear to be united. They open into the anterior, lobed part of the seminal vesicle, which has a flattened epithelium. The seminal vesicle may contain eosinophilic material, possibly related to the formation of a spermatophore. There is a large valve-like diaphragm separating the seminal vesicle from the ejaculatory duct. The opening of the ejaculatory duct is terminal but the tip of the penis papilla is bent downwards so that the opening appears ventral. The ejaculatory duct appeared widened distally in two specimens where a spermatophore was present. The penis papilla is large, blunt and cylindrical. It is fairly muscular with outer, subepithelial circular and inner longitudinal muscle layers. Eosinophilic tissue interspersed with longitudinal muscle fibres lies internal to the longitudinal muscle layer (Figs 10, 11).</p> <p>The adenodactyl is of the hollow type with a distinct lumen in the distal part of the papilla. The bulb may be over 250 µm in diameter and is muscular only at the most anterior part, and the muscle fibres mingle with those of the adjacent penis bulb. A central core of darkly staining cyanophilic tissue, interspersed with longitudinal muscle fibres, extends from the centre of the bulb throughout the length of the papilla, surrounding the duct in the distal part. The papilla may reach a length of 600 µm. Beneath the outer epithelium of the anterior part of the papilla there is a thick layer of circular muscle. This is underlain by a thinner layer of longitudinal/diagonal muscle which surrounds the cyanophilous glandular core. In the central part of the papilla, anterior to the lumen, the longitudinal/diagonal muscle layer becomes thinner and is absent from the muscle coat in the region of the lumen. In the posterior part of the papilla, there is a thin outer layer of circular muscle under which lies the glandular core and in the centre of this is the adenodactyl lumen. The lumen is surrounded by a cuboidal epithelium with insunk nuclei, surrounded by a thin layer of circular muscle. The lumen of the duct is filled with a secretion that is usually cyanophilic but may be eosinophilic or a mixture of both.</p> <p>The male and common atria are usually distinct, although sometimes the tip of the adenodactyl papilla extends into the entrance of the bursal canal. The male atrium is lined by a flattened epithelium and a short gonoduct leads from it to the gonopore.</p> <p>Karyology. A total of 43 metaphase plates of D. salina were examined from two specimens each from En Sheva and En Soda and revealed a mitotic chromosome number of 16 (Table 6, Fig. 12). In addition, 34 metaphase plates from 6 specimens from the River Jordan also revealed a mitotic chromosome number of 16. All chromosomes were metacentric, and of these, a few were metacentric heterobrachial bordering on metacentric isobrachial. The chromosomes decreased gradually in size. Pair number 8 was in all cases a little less than half the size of pair number 1.</p> <p>Discussion. Most species of Dugesia which have been examined karyologically have a haploid complement of n = 8, but six are known with n = 9 and three are known with n = 7. Other populations may be markedly aneuploid (e.g. Pala et al. 1982). The species of Dugesia from Israel that have been examined karyologically are the widespread, fissioning D. sicula (formerly known in Israel as D. biblica), which is normally triploid, 3n = 27, + 1-5 supernumeraries, and another ecotype of D. sicula, a normally sexually reproducing diploid form, 2n = 18, which is distributed in the central hilly region of Israel. Another sexually reproducing species, D. golanica, known only from the cold spring-fed waters of Dan and Banias in the very north of Israel, and previously thought to be D. cretica (Meixner, 1928) (Bromley &amp; Benazzi 1991) was also examined karyologically, 2n = 16 (Bromley 1974). Thus, regarding karyotypes, D. salina is similar to most other species of Dugesia, including D. golanica, in having a haploid number of n = 8.</p> <p>Regarding the copulatory apparatus, there are only a few species of Dugesia which have a large adenodactyl. Adenodactyls sensu stricto, which have no known function, may vary in number, size and position in relation to the penis papilla, but are defined as strongly muscularized, glandular-parenchymatic folds of the copulatory apparatus and provided with a lumen (Stocchino et al. 2017). The large, well-developed adenodactyl with a lumen, seen in ex-fissiparous specimens of D. salina, is most similar to that of D. cretica, which also has a single large hollow adenodactyl. However, the adenodactyl of D. cretica described from Crete by De Vries (1984) arises ventrally from the base of the penis papilla, and is positioned ventrally to the penis, whereas the large adenodactyl of D. salina is situated to the right of the penis. In contrast to this, Kenk (1930) described the large adenodactyl of D. cretica (from Crete) as being situated ventrally and to the right of the penis papilla. De Vries (1988) studied material attributed to D. cf. cretica from Cyprus, which had a dorsally situated adenodactyl, while material from the Greek island of Sérifos had an adenodactyl arising laterally to the right of the penis papilla, in a position similar to that of D. salina. However, De Vries (1988) noted that the position of the adenodactyl of D. cretica apparently varies between the geographically isolated populations in the eastern Mediterranean. True adenodactyls sensu stricto, strongly musculo-glandular organs of the type seen in D. salina and D. cretica, are not common among members of the genus Dugesia. They have also been recorded in Dugesia bactriana De Beauchamp, 1959, which occurs in Afghanistan and Pakistan, and in Dugesia transcaucasica (Livanov, 1951) from the Caucasus (Stocchino et al. 2017). D. bactriana has three small to medium-sized hollow adenodactyls all near the gonopore, two of which are posterior to the penis and one is ventral to it. D. transcaucasica has three medium-sized adenodactyls with a small lumen, one dorsal to the left, one dorsal to the right and one ventral to the right of the penis.</p> <p>A. En Soda population</p> <p>Other differences between D. salina and D. cretica lie in the head shape (D. cretica has much more pronounced auricles than D. salina) and penis shape, which is conical in D. cretica, but is large, cylindrical and somewhat blunt-ended in D. salina, and curves ventrally at its tip. De Vries (1988) noted that earlier records of D. cretica from Turkey, Iran and Israel are invalidated and that this species is known with certainty only from the Mediterranean islands of Crete, Tinos, Sérifos and Cyprus, and also from Romania.</p> <p>Although ex-fissiparous specimens of D. salina were obtained only from En Sheva, and on which the morphological description is based, the populations from En Sheva and En Soda appear to be identical. However, the population from the River Jordan showed slight differences with regard to pigmentation, karyotype and larger size. The lack of fission of the River Jordan animals during the winter may indicate that this population is a distinct ecotype recently derived from the En Sheva population and dispersed via the salt-water carrier. De Vries (1985) stated that generally, freshwater triclads have a very low resistance to salinity. The animals from these three sites obviously show an adaptation to living in water with a high salinity and it is notable that no other species of planarian was ever found in these habitats. It was suggested by Solà et al. (2015) that Dugesia specimens collected by them from another saline spring site in Israel, Einot Huga, which is close to En Soda, with a salinity of about 2000 mg /l Cl´, may perhaps also be assigned to D. salina on the basis of molecular studies.</p> <p>Stocchino et al. (2005) referring to the high level of endemicity of planarians of the genus Dugesia in the Mediterranean region, note that a process of allopatric speciation is strongly supported by their fragmented habitat, the present high endemicity values and the morphological divergence in copulatory traits. The unusual and highly saline habitats where Dugesia salina was found, as well as the differences in its copulatory apparatus from its morphologically closest congener (D. cretica) support the provisional observations of Whitehouse (1914) that ‘ Planaria salina ’ is a good species and apparently endemic to saline waters where other Dugesia species cannot thrive.</p> </div>	http://treatment.plazi.org/id/03E95D02FFFEFFBFC8F4F97F1D17FEEC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Bromley-Schnur, Heather J.	Bromley-Schnur, Heather J. (2021): The genus Phagocata Leidy (Platyhelminthes, Tricladida) in Israel, a new species of Phagocata from Lake Kinneret, and an emended description of Dugesia salina. Zootaxa 4969 (2): 293-317, DOI: https://doi.org/10.11646/zootaxa.4969.2.4
