PLANTEAMIENTOS MANIPULADORES
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4.1.1. Cis multidentatus species group: Cis chinensis
The Cis multidentatus species group was recently revised by SOUZA-GONÇALVES &LOPES -ANDRADE (2018). The specimens from the Caspian Forest can be identified following their identification key: They have single punctuation and single vestiture on the elytra (Fig. 22O–
Q), and are thus not Cis aldabranus, Cis chujoi, or Cis paraliacus; and they have a fovea on the 1st ventrite (a small one, Fig. 22J), and are thus not Cis limanicus. This leaves Cis chinensis and Cis multidentatus. The specimens from the Caspian Forest consistently lack microreticulation on the pronotum (Fig. 22A–F), the setae on pronotum and elytra are wide (squamate), and the lateral carinae of the pronotum are well visible from above; this clearly leads to Cis chinensis, for which the male genital characters also fit (base of tegmen without a small angulation at middle; Fig. 23A,B). However, the distinction between Cis chinensis and Cis mikagensis remains doubtful. LAWRENCE (1991) writes that Cis mikagensis (occurring in Japan) has narrower lateral margins of the pronotum than Cis chinensis. SOUZA-GONÇALVES
& LOPES-ANDRADE (2018), however, state that Cis mikagensis and Cis chinensis are potentially conspecific (the name mikagensis then having priority and being the valid name).
They could not clarify this point, as they did not study type material of Cis mikagensis, and the point can neither be clarified herein. This taxonomic distinction has some importance, as both Cis chinensis and Cis mikagensis are among the few pest species known in Ciidae.
The specimens from the Caspian Forest correspond with all the so far published descriptive data for Cis chinensis (mainly LAWRENCE 1991, 2016; LOPES-ANDRADE 2008; ROSE 2009;
SOUZA-GONÇALVES & LOPES-ANDRADE 2018), without contradiction. There is thus no indication that these specimens may represent a previously undescribed species of the Cis multidentatus species group. It is also noted that in the Cis chinensis specimens from the Caspian Forest there was no intra-specific variation in the way that characters form a transition to Cis multidentatus or other species of this species group. There was neither any regular variation between specimens from different localities or different host fungi. Yet, C.
chinensis does generally show some variation in colour, shape of pronotum, and shape of anterocephalic projections, as evident from comparing figs. 11‒19 (specimens from South Africa) and figs. 20‒28 (specimens from Saipan on the Mariana Islands) in SOUZA -GONÇALVES & LOPES-ANDRADE (2018). Such variation could partly result from the use of different food sources (see LOHSE &REIBNITZ 1991 and section 4.3.1.).
LOPES-ANDRADE (2008) already reported a wide range of variation of male-specific characters in males of his Brazilian Cis chinensis, regarding the overall size of the projections, and whether the apex is acute or more rounded. He also reported males almost devoid of pronotal projections and with the anterior pronotal margin just slightly emarginated, almost as in females. Based on specimens from France, ROSE (2009: fig. 2a,c) also showed that males vary in the extent to which male-specific characters are developed.
The many specimens from the Caspian Forest suggest that in the development of male-specific characters there is no continuous range, but there are likely two distinct categories, i.e. low males with poor development, and high males with strong development. This reminds of a similar situation in Cis pickeri (LOPES-ANDRADE et al. 2009). Yet, it cannot be excluded that in the populations in the Caspian Forest there are rare specimens showing transitions between high and low categories of males, and that future studies will reveal more fluent transition in other populations of Cis chinensis. It is also noted that low males from the Caspian Forest were always distinguishable from females with regard to the shape of their
110 pronotum and anterocephalic projections.
4.1.2. Cis boleti species group: Cis submicans
The Cis boleti species group is quite difficult with regard to the outlining of its species, mainly because revisionary taxonomic work is missing. One difficulty lies in whether the often subtle postulated differences between “species” are truly constant differences between units that represent species, or might in part rather represent intraspecific variation, perhaps with a less constant distribution over specimens. A clear-cut taxonomic structure of the Cis boleti species group requires a broad comparative study of morphological characteristics (especially male genitalia) and selected DNA sequences across the critical members of this species group and across their entire distribution ranges.
In recent decades some of the previously distinguished “species” were claimed to be conspecific and their names to be synomyms; for instance, Cis setiger Mellié, 1848 has been synonymised with Cis villosulus Marsham, 1802 (see JELINEK 2008). In some cases the same species was described several times by different authors, based on populations from Europe, Asia, and North America. For instance, LOPES-ANDRADE et al. (2016) compared the male genitalia of Cis pistoria from North America and Cis submicans from Iran (specimens from present project) and Poland and found no clear differences; this suggests them to be conspecific. Yet, species from the Cis boleti group usually show fairly distinct differences in the male genitalia, especially in the shape of the apical parts of tegmen and penis; this is true, for instance, for Cis submicans, Cis villosulus, Cis rugulosus, Cis micans, and Cis boleti. Of several species, or of putatively (according to published identification keys) conspecific specimens from different regions, the male genitalia were studied herein for comparison (Fig.
29), though only for very few specimens each. The specimens from two disjunct localities in the Caspian Forest (Fig. 29A,B, in Mazandaran and Golistan) showed the same genital characteristics as a specimen from Poland (Fig. 29D) that had previously been competently identified as Cis submicans; the genitalia also agree with those shown for Cis submicans in LOHSE (1967, as “Cis micans”): tegmen with slightly constricted midlength part, apically with distinct but only moderately long lateral lobes and a notched median lobe. Cis villosulus (Fig. 29F; LOHSE 1967, as “Cis setiger”) differs from this by the much longer (and wider) lateral lobes of the tegmen. Cis boleti (Fig. 29G) also has much larger lateral lobes of the tegmen. In contrast, Cis rugulosus (Fig. 29C) and Cis micans (Fig. 29E) differ by the vestigial condition of the lateral lobes in both the tegmen and penis, the latter also having a very wide median lobe, which is rounded (almost semicircular) in Cis rugulosus and triangular in Cis micans.
While the identification of many specimens from the Caspian Forest as Cis submicans is clear-cut, the samples are likely to include further species of the Cis boleti species group, as suggested by the considerable variation in body characters that are used to distinguish species from this group: degree of transverse curvature and relative width of pronotum, visibility of pronotal lateral carinae from above, setation of these carinae, (un)evenness of pronotum (presence, degree and distribution of bulges and moulds on pronotum), details of punctuation and setation of pronotum and elytra, body teint and shape characteristics, e.g. steepness of posterior part of elytra, and relative size of male abdominal fovea (see identification key of A. LOMPE at http://coleonet.de/coleo/texte/cis.htm). This topic could not be explored in the present work.
4.1.3. Cis comptus species group: Cis comptus and Cis striatulus
111 The Cis comptus species group has been intensely studied in recent years (LOPES-ANDRADE
et al. 2003; LOPES-ANDRADE 2004). The group includes the following species: Cis comptus, Cis striatulus (the two species found in the Caspian Forest), Cis clavicornis Baudi, 1873 (Cyprus, Turkey), Cis seriatocribratus Reitter, 1913 (Central Asia), Cis orius Kompantsev, 1996 (Central Asia, northern India), Cis tauriensis Królik, 2002 (Turkey: Taurus Mountains), Cis nepalensis Dodelin, 2011 (Nepal: Langtang Khola) from the Palaearctic region; Cis striolatus Casey, 1898 and Cis versicolor Casey, 1898 from the Nearctic region; and Cis fiuzai Almeida & Lopes-Andrade, 2004 (Brazil: Minas Gerais), Cis gumiercostai Almeida &
Lopes-Andrade, 2004 (Brazil: Minas Gerais), and Cis leoi Lopes-Andrade, Gumier-Costa &
Zacaro, 2003 (southeastern Brazil) from the Neotropical region. Among these, Cis nepalensis and Cis tauriensis have been suspected to be conspecific with Cis comptus (C. Lopes-Andrade, personal communication), and the Nearctic Cis striolatus is most likely conspecific with Cis striatulus (suspected by LAWRENCE 1971; synonymisation by LOPES-ANDRADE et al.
2016). Important characters of the group are the dual, more or less seriate elytral punctuation and the lack of conspicuous anterocephalic projections in males. The species of the group are partly very similar, but there are usually distinct differences in the male genitalia.
Regarding the samples from the Caspian Forest, Cis clavicornis can be excluded, as it has an antennal clava only comprising two clavomeres (KOMPANTSEV 1996). Cis orius can be excluded by its different male genitalia and its differently shaped sternite VIII. Cis comptus and Cis tauriensis (and Cis orius) have a stouter body than Cis striatulus and Cis seriatocribratus. Cis striatulus and Cis seriatocribratus differ in the length ratio between the 3rd and 4th antennomeres (= 1st and 2nd funiculomeres): 3rd much longer than fourth in Cis striatulus, and length subeaqual in Cis seriatocribratus. Cis comptus, Cis tauriensis, and Cis nepalensis have very similar male genitalia and might be conspecific; yet, Cis tauriensis shows a more strongly sclerotised penis tip at midline. By these criteria, the identification of Cis comptus and Cis striatulus among the Palaearctic members of the species group is clearcut ‒ under the premise that Cis comptus, Cis tauriensis, and Cis nepalensis represent the same species: Cis comptus.
Considering the two species of the Caspian Forest alone, Cis striatulus is more slender, its setae are finer, and the punctuation is more regular than in Cis comptus. However, in some cases dissection of genitalia was required to identify specimens as the one or the other species. Cis comptus has a short aedeagus and a tegmen similar to Ennearthron cornutum;
Cis striatulus has an elongate aedeagus and a very different tegmen.
4.1.4. Cis punctulatus species group: Cis tomentosus and Cis reitteri
Cis reitteri (synonym: Cis setifer Reitter, 1884) and Cis tomentosus are together with Cis punctulatus Gyllenhal, 1827 (European) and perhaps Cis libanicus Abeille de Perrin, 1874 (Lebanon only) the Palaearctic members of the Cis punctulatus species group. The Nearctic Cis horridulus Casey and Cis hystriculus Casey add to this.
Cis reitteri and Cis tomentosus are in many aspects very similar. In particular, their male genitalia are not distinguishable (Figs. 41A–D, 45A,B, Y), the abdominal fovea is completely absent (in both sexes). In Cis reitteri the pronotum is anterolaterally more projected and the lateral edge is more visible from above. The pronotal and elytral punctuation is dual, more distinct, and seriate in Cis reitteri, whereas punctuation in Cis tomentosus is single and hardly seriate, with punctures smaller and closer to each other than in Cis reitteri. Setae on the elytra are longer, more seriate, and more elucidate in Cis reitteri. Tibia denticle longer and more robust. Anterocephalic projections are thicker and larger in Cis reitteri. There are differences in the punctuation of the elytra.
112 In the Caspian Forest, Cis reitteri was always found in a low number in populations of Cis tomentosus, and their genitalia are not distinguishable. Their relationship is thus similar as that between Cis rugulosus and Cis villosulus (see section 4.1.2.), and there is some suspicion that the two might be conspecific, i.e. two morphs of the same species. While this should be kept in mind as a possibility, they are treated here separately, because they are morphologically distinctive.
REITTER (1901) introduced a variety of Cis tomentosus as Cis tomentosus var. tomentoides, from the Caucasus; it is listed as a synonym of Cis tomentosus in JELINEK (2008). As this variety occurs in the Caucasus, the point is of interest. Cis tomentosus var. tomentoides has been considered to differ from the nominate form as follows (REITTER 1901): it is matt; the vestiture is yellow, often brown-yellow in mature adults, the pronotum has a cloudy vestiture and a simple anterior edge; the anterocephalic edge forms two unclear humps less wide than in Cis tomentosus. The nominate form of Cis tomentosus is shiny, the vestiture is brown to black (yellowish brown only in immatures), the pronotum has no cloudy vestiture and a thickened anterior edge; the anterocephalic edge forms two obvious humps wider than in Cis tomentosus var. tomentoides. In the sampling for this project no specimens complying with the specific features of Cis tomentosus var. tomentoides were found, only Cis tomentosus in the strict sense.
4.1.5. Cis castaneus species group: Cis castaneus and Cis lugowoji
The Cis castaneus species group includes Cis castaneus, Cis lugowoji (the two species clearly found in the Caspian Forest during the current project), Cis lineatocribratus Mellié, 1848, Cis jacquemartii Mellie, 1848 (Europe), Cis matchanus Reitter, 1915 (Europe, Azerbaijan), Cis lasoni Królik, 2016, Cis glabratus Mellié, 1848 (Europe), Cis nikkoensis Nobuchi, 1960 (Japan), Cis yamamotoi Kawanabe, 1997 (Japan), Cis capricornis Kawanabe, 1997 (Japan), Cis laevigatus Kawanabe, 1997 (Japan), Cis konoi Chûjô, 1940 (Japan, Sakhalin), Cis rufocastaneus Nakane & Nobuchi, 1955 (Japan: Honshu), and, as the only Nearctic species, Cis levettei Casey, 1898 (North America). Members of this group were previously outlined as a subgenus of Cis: Eridaulus Thomson, 1863 (not Eridaulus Thomson, 1859, which refers to the Cis comptus group as valid designation; four years later Eridaulus Thomson, 1863 refers to the Cis castaneus group, synonym Cis nitidus group; JELÍNEK 2007).
The many Japanese species and the one from North America were not considered in the identification of the specimens from the Caspian Forest. Cis castaneus was identified by the anteriorly projecting anterolateral corners of the pronotum and by the presence of lateral hooks on the median lobe of the tegmen (see identification key of A. LOMPE at http://coleonet.de/coleo/texte/cis.htm). Cis lugowoji, which was often found to co-occur with Cis castaneus in basidiomes, lacks these distinct projections, like the other Eurasian species in this group.
KRÓLIK (2016) discussed the case of the names lineatocribratus, matchanus, and hanseni (the following text is modified from KRÓLIK 2016): REITTER (1915) described Cis lineatocribratus var. matchanus Reitter, 1915, as a variety of Cis lineatocribratus Mellié, 1849 based on specimens from the Balkan and Azerbaijan (Talysh Mts). According to REITTER (1915), Cis lineatocribratus var. matchanus differs from the typical form in the sculpture of the elytra: In the typical form the rows of punctures are coarse and deep, and the interspaces have only a few, barely visible punctures (thus appearing quite smooth). In var.
matchanus the rows of punctures are somewhat less strongly and deeply pronounced, and the interspaces have dense and fine punctures. This makes the elytral sculpture of var. matchanus more similar to that in Cis jacquemartii and Cis castaneus, in which the coarser elytral punctures, however, do not form distinct rows. The two teeth of the anterocephalic edge
113 (clypeal teeth) are sharper in var. matchanus than in the typical form [end of citing information from KRÓLIK 2016]. KRÓLIK’s (2016) studies of external morphology and male genitalia further led him to the conclusions that var. matchanus Reitter, 1915 is (1) not conspecific with lineatocribratus Mellié, 1849, and is (2) conspecific with hanseni Strand, 1965. Following the priority of the earlier name matchanus versus the later hanseni, this was reason to accept Cis matchanus as the valid name for the species so far called Cis hanseni (which is now a synonym of Cis matchanus).
The very similar Eurasian species Cis lugowoji, Cis lasoni, Cis matchanus, Cis glabratus, and Cis lineatocribratus have been separated in the literature by characters concerning the presence of (always small) projections on the male anterocephalic edge, the shape of the pronotum and the visibility of its lateral carinae (both characters with pronotum seen from above), the difference between and density of the two types of elytral punctures, the degree of seriality of elytral punctures (especially near the suture), and the shape of tegminal and penial parts of the male genitalia. The samples from the Caspian Forest show much variation in these characters. This variation could not be analysed in the frame of the present thesis. Some specimens could be clearly identified as Cis lugowoji based on the shape of the male genitalia, but other specimens showed somewhat different genital features. The samples from the Caspian Forest thus likely included a mixture of Cis lugowoji and one or more of the abovelisted similar Eurasian species. More research is needed, especially the comparative study of genitalia and molecular data (e.g. DNA-barcoding) for numerous specimens of the samples from the Caspian Forest.
4.1.6. Cis fissicollis (not assigned to a species group)
Cis fissicollis is quite distinct from other Palaearctic Ciidae species by its distinct median longitudinal groove on the pronotum. While Cis fissicollis and Cis fissicornis Mellié, 1848 are not assigned to a species group, the two are quite similar. The morphology of the male genitalia has not been described for Cis fissicornis, so that a comparison of these is not possible. However, Cis fissicollis can be distinguished by (nearly) the entire lateral margins of the pronotum are visible from above.
As in Cis chinensis (see section 4.1.1.), male-specific features are found in two distinct degrees in Cis fissicollis, i.e. there are low males and high males. In the Caspian Forest, males were distinctly rarer than females in nearly all collected samples (see Appendix 9.1.);
the summary ratio was 50 males versus 107 females.
4.1.7. Cis festivus species group: Cis festivus
The Cis festivus group includes Cis festivus (found in the Caspian Forest), Cis vestitus Mellié, 1848, and Cis pygmeus Marsham, 1802 from the Palaearctic region, and some Nearctic species, among which Cis festivulus is very similar to Cis festivus. For some time, members of the Cis festivus group were assigned to the genus Orthocis because of the similar rounded condition of the apical dorsal edge of the tibia; but the group was returned to the genus Cis (LAWRENCE 1971) because the not elongated 3rd antennomere and the distinct anterocephalic projections of the male are in conflict with the defining criteria of Orthocis.
The fringe of raised and reflexed setae on the anterior edge of pronotum is in Cis festivus longer than the lateral fringe, whereas in Cis vestitus it is shorter than the lateral fringe, often virtually absent. The lateral carina of the pronotum is in Cis festivus more expanded and posteriorly wider than in Cis vestitus. The strong reticulation on the pronotum points to Cis vestitus, while weak reticulation is ambiguous. The antennal club is pale or brownish in Cis festivus but dark and blackish in Cis vestitus. In Cis pygmeus the lateral edge of the pronotum
114 is narrower, and it has a vestiture with golden-brown setae; setae are moderately fine and of unequal size; the pronotum is widest posteriorly and anteriorly narrower, and has a punctuation denser than on the elytra. These characters allowed for the specimens from the Caspian Forest a clear identification as Cis festivus.
4.1.8. Genus Ennearthron: Ennearthron cornutum
Ennearthron includes 14 species from the Palaearctic region, 6 of them only known from Japan, and 1 from there and nearby China (JELINEK 2008). A few additional species are known from other zoogeographical regions. The specimens from the Caspian Forest were identified to species as follows: Ennearthron reitteri has a short cylindrical, robust body (similar to Cis comptus), a densely punctured pronotum (indistinctly seriate), short vestiture with setae of unequal size, the lateral carina is visible from above and anteriorly narrower. In Ennearthron pruinosulum the lateral edge of pronotum is visible from above, finely wrinkled and uneven, the vestiture is dusty and very short, elytral length/width ratio (EL/EW) is 2.5.
Ennearthron filum is a very small and narrow species (0.5–1.1 mm), the vestiture is short and whitish. Punctuation of pronotum and elytra of Ennearthron palmi is moderately fine and rather irregular. These characters allowed for the specimens from the Caspian Forest a clear identification as Ennearthron cornutum.
The population of Ennearthron cornutum from the Caspian Forest is very special compared to other Palaearctic populations (as far as these have been examined for this character) by the complete lack of the abdominal fovea in the male. I decided not to take this as the basis for the description of a new species, as in all other characters the Caspian population fully agrees with other descriptions for Ennearthron cornutum, and because there is another case of likely
The population of Ennearthron cornutum from the Caspian Forest is very special compared to other Palaearctic populations (as far as these have been examined for this character) by the complete lack of the abdominal fovea in the male. I decided not to take this as the basis for the description of a new species, as in all other characters the Caspian population fully agrees with other descriptions for Ennearthron cornutum, and because there is another case of likely