Chordodes undulatus Linstow, 1906

Schmidt-Rhaesa, Andreas, 2002, Australian species of Chordodes (Nematomorpha) with a description of two new species, remarks on the genus Chordodes and its life history, Journal of Natural History 36 (13), pp. 1569-1588 : 1573-1584

Chordodes undulatus Linstow, 1906

Chordodes undulatus Linstow, 1906b: 257–258 , gure 20.

HOLOTYPE. Sydney , Australia, one female (Linstow, 1906b).

Material examined. None.

Host. An undetermined praying mantid.

This species is very similar to ‘ Chordodes ’ annulatus . It also lacks crowned areoles and therefore does not belong to the genus Chordodes . Instead, two types of areoles are present equivalent to ‘ Chordodes ’ annulatus . The only diVerence is that in ‘ Chordodes ’ undulatus a tubercle may be present on some of the large areoles.

Chordodes queenslandi nov. sp.

(gures 3–6)

Chordodes bouvieri: Römer, 1895: 797–799 .

2014 Chordodes modiglianii: Heinze 1935: 24 .

HOLOTYPE. Queensland, Australia: Carnarvon Gorge National Park , Moss February H Garden Entomology. and A and. Zwick) Hell in Canberra. Hole Deposited Gorge , Australia at (25 the ss00, Australian ¾S accession, 148ss00¾E number National); female 9 Insect 500; 13 002 March Collection 232. 1997; (coll CSIRO. P., 13 PARATYPE. Queensland, Australia: Carnarvon Gorge National Park , Moss 39: Garden and Hell Hole Gorge (25ss00¾S, 148ss00¾E); male; 13 March 1997; coll. P.,

23 H. and A. Zwick. Deposited at the Australian National Insect Collection ( CSIRO at Entomology) in Canberra, Australia, accession number 9 500 002 233.

]

Bath Dämel Further (Römer record, 1895. Sydney), same, Australia specimen, one as female C. modiglianii ; as C. bouvieri (Camerano (Villot,, 1885 1892)), coll (see. of Heinze, 1935) (Zoologisches Museum der Universität Hamburg, Germany; V 2231). University (Mantoptera Host. Holotype). and paratype: female of Sphodropoda tristis (Saussure, 1871) [Description of female

by The large female is 20.5 cm long and 1.2 mm in diameter. The body is tightly Downloaded The body tapered lled body with (, gure the colour oocytes cloacal 3A, is B) as a. opening light The can ventral brown be is seen situated with side in cross-sections numerous is terminally marked darker by on. The a the very brown posterior anterior darkly patches end end pigmented is (all gure distinctly over 4 line F the). (gure 3A, C). These diVerent coloration patterns are not re ected in diVerences of the surface structures.

The whole body is covered with protruding structures called areoles. The cuticle between the areoles is structured into cord-like folds (gure 4E). Six types of areole can be distinguished which are numbered consecutively in the following for better orientation. Most abundant are more or less circular or oval areoles (type 1; gures 3B, E, 4A, B) with diameters of 5–10 m m. Among these are scattered areoles of the same shape but with a tubercle on top (type 2; gures 3B, E, 4A, B, D). These tubercles often originate in a slight depression on top of the areole. They are about 10 m m long and do not taper towards the tip. The rarest type of areole has a solid spine on top (type 3; gures 3E, 4G). This spine originates, like the tubercles, in a slight depression and is slightly longer than the tubercles. The spines have not been found in the posterior end of the animal, but are scattered in the anterior and middle region.

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The three other areolar types are more elevated than the rst three and occur in typical clusters. They are always darker than the previous types. Within dark patches they appear as very dark brown structures and within the light brown cuticular regions they are coloured a medium brown. Areoles of type 4 occur abundantly in clusters of two, three or more over the whole cuticle (gures 3B, E, 4A, B). They are approximately three to four times higher than the surrounding areoles of types 1–3. The apical surface is structured with numerous wart-like structures that can even form very short and stout processes. The two remaining areolar types both 2014

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2014 February 13 39: 23 at] Bath of University [by Downloaded have laments on top and are therefore called crowned areoles. Crowned areoles occur with very long laments (type 5) and with moderately short laments (type 6). Both types of crowned areoles always occur in clusters of two and are surrounde d by about 8–12 elevated areoles of type 4. The crowned areoles with long laments only occur on both sides along the ventral and the dorsal midline (gures 3A, C, D, 4G). The clusters are 80–120 m m apart from each other. They do not have equal distances from the midline or from each other, but are only roughly arranged in two lines on either side of the midline. The number of laments per areole varies and obviously the laments can break oV leaving only short rudiments. The laments are about 200 m m long. The crowned areoles with shorter laments (type 6) are distributed all over the cuticle (gures 3B, E, 4A–C). They also occur in clusters of two surrounded by elevated areoles. The two clustered crowned areoles enclose a central tubercle between them (gures 4A, C). The existence of a central tubercle

2014 February 13 39: 23 at] Bath of University [by Downloaded could not be observed with the long lamented areoles (type 5) but was not clearly excluded either. The apical surface of the crowned areoles is more or less at with ridge-like structures that give rise to the laments on the lateral side (gure 4C). Most laments are 5–15 m m long, but can also reach lengths of about 25 m m.

Areoles cover the whole body except for the anteriormost and posteriormost ends. The region around the cloacal opening is free of areoles (gure 4F). Scattered bristles are present between areoles in a ring-like region around the cloacal opening (gure 4F). The elevated areolar types are not present close to the posterior end, the areolation consists of types 1 and 2.

Description of male

The male is 15.5 cm long with a maximum diameter of 1 mm. It is slightly attened, probably due to already-released gametes. The anterior end is distinctly tapered. The body colour is darker than in the female, but the same pattern of dark patches is present. A ventral midline is indicated by the pattern of dark patches, but is not as clearly marked as in the female (gure 5A).

The cuticle contains the same six types of areole as the female, but in slightly diVerent patterns. In contrast to the female, the cuticle between the areoles is smooth and not structured into ridges (gure 5C). Areoles stand very close together in the anterior end, but slightly further away from each other in the middle region of the body. Areoles of type 1 are abundant, but appear in diVerent sizes which probably represent transitions to type 4 areoles (gure 5C). Many type 1 areoles are structured apically similar to type 4 (gure 5B, C). Areoles with tubercles (type 2, 5tubercle areoles) seem to be more elevated than in the female and less abundant (gure 5B, C). This type is very scarce in the anterior region of the body and slightly more abundant in the median and posterior region. The thick spines of type 3 areoles are more abundant and conspicuous than in the female (gure 5B, C). Apart from possible transitions between types 1 and 4, fully diVerentiated type 4 areoles can be found in similar shape and distribution as in the female. They appear individually or in clusters of two. Clusters of 8–10 of these areoles always surround areoles of types 5 and 6 (gure 5B–D). Crowned areoles of both types occur in the male and areoles with long laments (type 5) are distributed in paired rows along the ventral and dorsal midline (gure 5A, D, E). Areoles with short laments (type 6) are present all over the cuticle (gure 5B, C, H). Both types of crowned areoles are always clustered in groups of two, but a tubercle between these two areoles could not be observed. From a lateral view areoles of type 6 appear to contain a canal leading from the base to the apical surface (gure 5F, G).

The cloacal opening is situated ventrally, at about 270 m m distance from the posterior margin of the animal. Fine structural details of the cloacal opening could not be observed optimally, but it appears to have an oval shape and no associated structures such as circumcloacal spines which are present in several other nematomorph taxa. Anterior of the cloacal opening is a ventral strip of 50–70 m m width structured only by very at areoles (gure 6B). At the lateral margins of this strip, areoles of type 2 are distributed, bordered by the usual areolation (gure 6B). A few ne bristles are situated anterolaterally of the cloacal opening (gure 6A, small arrows). Posterior of the cloaca there is a paired oval region that is slightly infolded (gure 6A). The margin of this region is structured by at hexagonal or rounded areoles that are bordered in the lateral and posterior region by more elevated areoles. Laterally of the cloacal opening is a paired oval region of about 80 m m width and

145 m m length bearing numerous bristles (gure 6A, large arrows). These bristles become gradually sparser towards the margins of this bristle eld and intermingle with the surrounding areoles (gure 6D). There are also abundant minute bristles (gure 6D), most of which occur on the areoles, sometimes like a crest (gure 6C). These minute bristles are very abundant on areoles at the posterior tip of the animal and more scarce on areoles lateral of the bristle elds.

Specimen Zoologisches Museum Hamburg V 2231

Römer (1895, 1896) described a specimen collected in Sydney, Australia and assigned it to Chordodes bouvieri (Villot, 1885) which otherwise has been reported only from South America. After a re-examination, Heinze (1935) assigned this specimen to the South-East Asian species C. modiglianii (Camerano, 1892) . A further reinvestigation of this specimen showed that areolar types 1, 2, 4, 5 and 6 (as described above for C. queenslandi ) (gure 6E

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and that in particular 2014 the areoles of type 5 occur in paired rows along the ventral and dorsal longitudinal

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line The 1915.) ventral This. Areoles pattern line of is type is not neither 3 as could marked present not be by in observed C a. dark bouvieri , pigmentation but nor this in may C., modiglianii be but due the to body their (Camerano scarcity shows a., pattern of dark pigmented areas comparable to the one of the two specimens

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Bath HOLOTYPE. Queensland, Australia: Goldsborough State Forest Reserve (17ss14¾S,

of 145ss47¾E); male; 28 March 1997; coll. P., H. and A. Zwick. Deposited at the

University accession

Australian Host. Male number National of Hierodula 9 500 Insect 018 Collection 479 majuscula . ( CSIRO (Tindale Entomology, 1923) (Mantoptera) in Canberra)., Australia,

[The specimen is 6.5 cm long with an approximate diameter of 0.7 mm. The

by diameter is diYcult to measure exactly because the specimen is strongly attened.

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The colour clusters The anterior is, cuticle between dark end brown contains which is tapered, without regions diVerent, a patches white free types of tip such areoles of or areoles as dark are in C which collar visible. queenslandi are are (gure distributed not. 7 present B). These. in The irregular regions body are structured by regular, cord-like folds (gure 7A). There are three types of areoles which are distinctly elevated. All have diameters between 11 and 14 m m. Crowned areoles have extremely short apical laments (gure 7A–D). They appear as paired clusters with an intermediate tubercle (gure 7A, C). There is no second type of crowned areole as in C. queenslandi and the ventral and dorsal midline are not distinctly bordered by lines of crowned areoles. The clusters of crowned areoles are not surrounded by type 4 areoles (as in C. queenslandi ) (gure 7A–D). Crowned areoles sometimes appear as single structures (gure 7A, arrow) with irregular distribution over the cuticle. In these areoles the apical laments are extremely short and sometimes diYcult to recognize. When this is the case, the areoles resemble the type 4 areoles of C. queenslandi and a transition between type 4 areoles and crowned areoles cannot be excluded. The third elevated areolar type are tubercle areoles (type 2; gure 7A, B). They are comparable in size to the other elevated types. The tubercle is a distinct, nger-like projection. It appears as if it originates from a depression in

2014 February 13 39: 23 at] Bath of University [by Downloaded the areole which therefore sometimes appears to have a horseshoe-like rim (gure 7B, top) or even to be separated into two parts (gure 7A). Additionally, there are very small tubercle areoles distributed over the whole cuticle (gure 7A). These measure about 2.7 m m in diameter, are slightly elevated and have a short, delicate tubercle on top.

The cloacal opening is situated at about 200 m m distance from the posterior margin of the animal (gure 8A). It could not be observed if circumcloacal spines are present, because the cloacal opening is covered by a mass of spermatozoa. Anterolateral of the cloacal opening are paired regions in which numerous bristles are located (5bristle elds) (gure 8A). Posterior is a region devoid of areoles with only a few spines being present. The midline anterior of the cloacal opening is structurally diVerent from the remaining cuticle (gure 8A, B).

Within the bristle elds, there are long slender bristles, more solid spines and numerous minute bristles (gure 8C). Areoles are almost absent from the bristle elds; when present, they are hardly elevated. The spines may be modi ed tubercle areoles. In the marginal regions of the bristle eld the tubercle areoles have a more or less pointed tubercle (gure 8E) and it appears as if these ‘spine-like tubercles’ become separated by a progressive attening of the corresponding areoles.

The region with long bristles is on each side at a distance of about 35 m m from the cloacal opening and extends 70 m m in diameter and 140 m m in longitudinal direction.

The bristles are maximally 21 m m in length. At least the majority is unbranched

(gure 8C), but a few bristles appear to be branched apically. Interspersed between bristles and spines are minute bristles of less than 2 m m length (gure 8C). These bristles also appear in the marginal regions of the bristle elds where long bristles

2014 February 13 39: 23 at] Bath of University [by Downloaded are absent. They occur in the interareolar spaces as well as on the areoles themselves (gure 8E). These areoles with minute bristles are distributed lateral of the bristle elds and on the posterior tip of the animal, comparable to C. queenslandi .

The region posterior of the cloacal opening does not contain recognizable areoles, but it is structured by numerous deep grooves that create irregularly shaped compartments (gure 8D). Among these structures are, more or less radially around the cloacal opening, short spines which originate in the furrows (gure 8A, D). These spines extend on both sides of the cloacal opening to the ventral midline anterior of the cloacal opening. This midline is structurally diVerent from the lateral parts. It contains the cord-like structure of the cuticle as described above with irregularly formed at areoles (gure 8B). These areoles have a smooth surface. On the margins of the ventral midline there are also tubercle areoles (gure 8B). The tubercle is apically acuminated and spine-like. At lower magni cations, these spines on areoles appear as a continuation of the spines posterior of the cloacal opening. The whole ventral midline region is 75 m m broad.

Chordodes jandae Camerano, 1895 (gure 9) Chordodes jandae Camerano, 1895: 8–9 .

Type series. Timor , Indonesia: stream close to Dillu; two males; coll. H. ten Kate (Camerano, 1895) (National Museum of Natural History Naturalis, Leiden, The Netherlands; RMNH Vermes 838) .

Further specimens. Queensland, Cucania, Australia: female; coll. W. E. Schevill, April 1932 ( Museum of Comparative Zoology , Harvard University, MA, USA; MCZ 2093 ) .

Material examined. Specimen from Australia ( MCZ 2093), cuticular preparation (light microscopy) and SEM preparation from midbody.

Host. From ‘green mantid’ (on label).

The same specimen was investigated and described by Miralles and Villalobos (1994). Their results can be con rmed and are brought into context with the descriptions of C. queenslandi and C. brevipilus .

Chordodes jandae possesses areolar types 1–4 and 6 as described for C. queenslandi (gure 9A–C). They appear in comparable patterns. Crowned areoles (type 6) occur as pairs surrounding a central tubercle (gure 9A, B). They are surrounded by a circle of type 4 areoles (gure 9A–C). The apical laments of the crowned areoles are short (gure 9A). The solid spines (type 3, gure 9A–C) were not mentioned by Miralles and Villalobos (1994). Tubercle areoles (type 2) occur rarely (gure 9C).

Miralles and Villalobos (1994) described large pores or ‘perforated areoles’. Similar structures could be observed in the microscopical cuticular preparations. There they appear as large oval structures surrounded by type 4 areoles (gure 9D). It cannot be excluded that these are areas where structures such as the crowned areoles were lost, probably mechanically.

The original description of C. jandae (Camerano, 1895; gures in Camerano, 1897) does not indicate the presence of spines (type 3). Additionally, type 4 areoles appear to be slightly diVerent and have an irregularly structured surface (like a strawberry according to Camerano (1895)) in the specimens from Timor. If the

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6 2093. (D)), ‘ cuticle Crater’, SEM surrounded and light by areoles microscopy of type. (A 4 –. C (A)) Cuticle SEM,

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Downloaded Host further assignment –parasite reinvestigations of correlation the Sydney of specimen the type specimens to C. jandae . is correct remains to be shown by Both specimens of Chordodes queenslandi were observed to emerge from their host, adult females of the mantid Sphodropoda tristis (Saussure, 1871) by the collectors. The rst mantid fell into water during an escape from capture and C. queenslandi emerged immediately. A second specimen of S. tristis was captured, which showed no signs of parasitism. When its abdomen was brie y dipped into water, C. queenslandi emerged within 15 s. The behaviour of the hosts did not change after emergence of the parasites. The only diVerence was that the abdomen was completely hollow as could be observed by contact, but the internal anatomy was not further investigated. The same experiment (bringing captured mantids into contact with water) was repeated with two diVerent mantid species ( Kongobatha diademata Hebard, 1920 and Hierodula majuscula (Tindale, 1923)) in the Goldsborough State Forest Reserve ( SW Gordonvale, Kearney Falls) with the same result. One of the emerged nematomorphs is C. brevipilus , the other belongs to a yet undetermined species.

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