Widespread and persistent invasions of terrestrial habitats coincident with larval feeding behavior transitions during snail-killing fly evolution (Diptera: Sciomyzidae)

BACKGROUND: Transitions in habitats and feeding behaviors were fundamental to the diversification of life on Earth. There is ongoing debate regarding the typical directionality of transitions between aquatic and terrestrial habitats and the mechanisms responsible for the preponderance of terrestrial to aquatic transitions. Snail-killing flies (Diptera: Sciomyzidae) represent an excellent model system to study such transitions because their larvae display a range of feeding behaviors, being predators, parasitoids or saprophages of a variety of mollusks in freshwater, shoreline and dry terrestrial habitats. The remarkable genus Tetanocera (Tetanocerini) occupies five larval feeding groups and all of the habitat types mentioned above. This study has four principal objectives: (i) construct a robust estimate of phylogeny for Tetanocera and Tetanocerini, (ii) estimate the evolutionary transitions in larval feeding behaviors and habitats, (iii) test the monophyly of feeding groups and (iv) identify mechanisms underlying sciomyzid habitat and feeding behavior evolution. RESULTS: Bayesian inference and maximum likelihood analyses of molecular data provided strong support that the Sciomyzini, Tetanocerini and Tetanocera are monophyletic. However, the monophyly of many behavioral groupings was rejected via phylogenetic constraint analyses. We determined that (i) the ancestral sciomyzid lineage was terrestrial, (ii) there was a single terrestrial to aquatic habitat transition early in the evolution of the Tetanocerini and (iii) there were at least 10 independent aquatic to terrestrial habitat transitions and at least 15 feeding behavior transitions during tetanocerine phylogenesis. The ancestor of Tetanocera was aquatic with five lineages making independent transitions to terrestrial habitats and seven making independent transitions in feeding behaviors. CONCLUSIONS: The preponderance of aquatic to terrestrial transitions in sciomyzids goes against the trend generally observed across eukaryotes. Damp shoreline habitats are likely transitional where larvae can change habitat but still have similar prey available. Transitioning from aquatic to terrestrial habitats is likely easier than the reverse for sciomyzids because morphological characters associated with air-breathing while under the water\u27s surface are lost rather than gained, and sciomyzids originated and diversified during a general drying period in Earth\u27s history. Our results imply that any animal lineage having aquatic and terrestrial members, respiring the same way in both habitats and having the same type of food available in both habitats could show a similar pattern of multiple independent habitat transitions coincident with changes in behavioral and morphological traits

On taxonomy of the subgenus Amblycladius of the genus Chaetocladius (Diptera, Chironomidae)

Krasheninnikov, Andrey B., Przhiboro, Andrey A. (2022): On taxonomy of the subgenus Amblycladius of the genus Chaetocladius (Diptera, Chironomidae). Zootaxa 5168 (4): 494-500, DOI: https://doi.org/10.11646/zootaxa.5168.4.1

Jurochlus trivittatus Lukashevich & Przhiboro, 2012, sp. nov.

<i>Jurochlus trivittatus</i> sp. nov. <p>(Figs 1 E, 3A–E, 4, 5)</p> <p> <b>Type material.</b> <i>Holotype</i>: PIN 4270/2581 (abdominal segments II–IX of female pupa). <i>Paratype</i>: PIN 4270/2540 (abdominal segments V–IX of male pupa). SW Mongolia, Gobi-Altai Aymag, Shar Teg (outcrop 443/1), Upper Jurassic.</p> <p> <b>Etymology.</b> From the Latin <i>vitta</i> (band), in reference to the pattern of three longitudinal stripes on abdomen.</p> <p> <b>Description.</b> <i>Pupa of female.</i> Each segment except terminal one with three wide nearly parallel (broadly oval or subrectangular) concavities, median and two lateral ones, darker than surrounding areas. Anterior margins of tergites well-sclerotized; posterior margins darkened. Tergal ridge very strong on segment II, becoming progressive smaller to more posterior segments (up to segment VII). Some segments with fine, more or less longitudinal (fan-shaped) striation (most distinct on II and V; probably traces of muscles) posterior to ridge; some such lines extending to anterior margin of next segment. Lateral parts of segments II–VII with elongate concavities (probably adhesion marks). No distinct LS setae visible.</p> <p>Segment IX. Outer margin of anal lobe sinuate, distinctly convex in middle part and strongly concave in apical third. Outer margin finely serrate, serration clearly visible in middle third of margin, consisting of triangular stout spines with wide bases (ca. 40 spines visible); inner margin with spines of similar size and shape at least in its middle. Lobes apically tapered into dark spurs; distance between apical spurs 0.4x the width of segment IX at base. Genital sac slightly more pigmented than outer adjoining area, elongate, slightly tapering, with apically rounded parts, not reaching segment apex in its middle. Segment IX anterior to genital sac with darkened area at midline and at anterior margin.</p> <p> <i>Pupa of male.</i> Similar to female, but three concavities on segments not as distinct, dark pattern not visible. Tergal ridge strong on segment V, becoming progressively smaller on more posterior segments (up to segment VII). Tergite VIII with indistinct traces of possible ridge. Segments V and VI with fine, more or less longitudinal (fan-shaped) striation posterior to ridge.</p> <p>Segment IX with traces of two stout lateral setae in anterior third. Outer margin of anal lobe finely serrate, with triangular stout small spines at least in middle third; inner margin possibly with similar smaller spines (not distinctly visible). Genital sac with anterior parts more distinctly visible, darker than posterior ones, separated from these latter by diagonal lines. Areas anterior to genital sac well outlined laterally.</p> <p> <b>Measurements.</b> Female. Combined length of abdominal segments II–IX, 8000; entire body length (extrapolated), 10000–11000. Abdominal segments length: II–VIII, 900–1100; width: II–V, 2600; VI, 2500; VII, 2400; VIII, 2250. Abdominal segment IX: length, 1380; width at base, 2000; distance between tips of anal lobe spurs, 800.</p> <p>Male. Combined length of abdominal segments V–IX, 5190; entire body length (extrapolated), 10000–11000. Abdominal segments length: II–VII, 1050; VIII, 800; width: V–VI, 2380; VII, 2250; VIII, ca. 2130. Abdominal segment IX: length, 1380; width at base, 1870; distance between tips of anal lobe spurs, 750.</p> <p> <b>Note.</b> We suggest the male and female pupae to be conspecific based on the similar appearance and body proportions (especially that of segment IX), the presence of well-developed tergal ridges on the same segments, and the serration of anal lobe margins consisting of spines, which are similar in shape.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on page 439, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

Jurochlus Kalugina

Genus <i>Jurochlus</i> Kalugina <i>in</i> Kalugina <i>et</i> Kovalev, 1985 <p> <b>Type species</b> <i>Jurochlus sibiricus</i> Kalugina in Kalugina et Kovalev, 1985: 95, fig. 48.</p> <p> <b>Differential diagnosis (emended).</b> Medium-sized to large pupae, 4.5– 11 mm long. Thoracic horn widened distally, with well-developed plastron plate and stout basal stalk. All leg sheaths situated under wing sheath, meeting together at tip of wing sheath, distal portion of fore and midleg darkened sheaths practically straight, Sshaped portion of hind leg sheath beneath distal half of wing sheath. Segments II–VIII short and wide, nearly rectangular, with straight or slightly convex lateral margins. Anterior and posterior margins of abdominal segments nearly straight; anterior margins of tergites II–VIII appearing as sclerotized transverse stripes; posterior margins of tergites often pigmented; tergites II–VII usually with a well-developed transverse ridge in posterior part. Segment VII with two or three LS setae. Segment VIII with five LS setae in distal two-thirds, posterior angles not produced into large lobes (at most, with small, barely projecting lobes); female sternite VIII at posterior margin with accessory genital sacs separated by distinct longitudinal crease.</p> <p>Segment IX in proximal third with two lateral close-set setae on either side; setae short, about 0.2x the lobe length. Anal lobe more or less triangular, apically pointed into a dark spur. Outer and inner margins of anal lobes usually with continuous row of saw-like teeth; inner margins more finely serrate; serration on both margins not reaching spurs. Outer margin of anal lobe more or less straight or slightly sinuate (convex in proximal two-thirds and concave in apical part); inner margins converging, usually sinuate. Male genital sac wedge-shaped, with inner margin confluent, distally tapered, reaching 3/4 of lobe length and projecting outside anal lobe margin. Female genital sac shorter, slightly tapered, not projecting outside anal lobe margin; female genital plate IX transverse, distinct.</p> <p> <b>Species included.</b> Besides the type species, <i>J. rigor</i> Kalugina, 1985 from the same locality (Middle or Late Jurassic of Transbaikalia) and four species described below from the Late Jurassic and Early Cretaceous of Mongolia, all known from pupae only.</p> <p> <b>Remarks.</b> The genus was established based on two impressions of pupae attributed to two species. Up to now, only these two species have been known.</p> <p>Despite the abundance of impressions of adult chironomids at Shar Teg and Khutel Khara, and our examination of every impression from Shar Teg, no specimens of appropriately large size have been found, and thus no association can be made.</p> <p> Only the holotype of <i>J. sibiricus</i> is preserved completely, all other species are known only by abdomens. Hence, the structure of the thoracic horn, thorax and wing sheaths can be described only for the type species, known from a single impression.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on pages 435-436, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

Jurochlus adustus Lukashevich & Przhiboro, 2012, sp. nov.

<i>Jurochlus adustus</i> sp. nov. <p>(Figs 3 I, J, 8B, C)</p> <p> <b>Type material.</b> Holotype PIN 3965/3209 (female pupal abdominal segments III–IX). Mongolia, East Gobi Aymag, Khutel-Khara (=Hara Hutul), lower Tsagantsab Formation, Lowermost Cretaceous (Uppermost Jurassic not excluded).</p> <p> <b>Etymology.</b> From the Latin <i>adustus</i> (swarthy), in reference to dark pattern.</p> <p> <b>Description.</b> Female. Anterior margins of tergites appearing as wide slightly sinuate sclerotized stripes; posterior margins of all segments with a thin line, without traces of ridge. Posterolateral margins of tergites, as a rule, slightly projecting, rounded or bluntly tapered. Tergites uniformly moderately darkened except for lighter lateral margins; posterior margins slightly more darkened. LS setae and their attachment places not visible. Tergites III–VII with traces of short setae scattered over surface, most clear on segment V. Segments III–V posterior to hind margin of tergite (probably intersegmental membrane) with fine, more or less longitudinal (fan-shaped) striation (possibly, traces of muscles); some of such lines extending to posterior or anterior margins of tergites.</p> <p>Segment IX. Outer margins of anal lobes almost straight in apical two-thirds, slightly concave near apex, possibly indistinctly serrate in middle parts (no distinct teeth visible). Inner margins slightly sinuate. Anal lobes with strongly impressed stout spurs. Genital sac tapering, apically more or less truncate. Genital plate IX appearing as transverse stripe with margins slightly impressed.</p> <p> <b>Measurements.</b> Combined length of abdominal segments IV–IX, 5500; entire body length (extrapolated), 9500–10500. Abdominal segments length: IV–VII, 800; VIII, 870; width: III–VII, 2150–2200; VIII, 1920. Abdominal segment IX: length, 1250; width at base, 1500; distance between tips of anal lobe spurs, 500.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on page 446, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

Jurochlus lineatus Lukashevich & Przhiboro, 2012, sp. nov.

<i>Jurochlus lineatus</i> sp. nov. <p>(Figs 6, 7)</p> <p> <b>Type material.</b> <i>Holotype:</i> PIN 4270/2544 (part and counterpart of male pupal abdominal segments VII–IX). <i>Paratype</i>: PIN 4270/348 (part and counterpart of male pupal abdominal segments VII–IX). SW Mongolia, Gobi- Altai Aymag, Shar Teg (outcrop 443/1), Upper Jurassic.</p> <p> <b>Etymology.</b> From the Latin <i>lineatus</i> (rectilinear), in reference to the nearly straight outer margin of the anal lobe.</p> <p> <b>FIGURE 5.</b> <i>Jurochlus trivittatus</i> <b>sp. nov.</b> from Shar Teg, J3: A, B, holotype PIN 4270/2581, pupa of female: A, segments IV–IX, B, segments VIII and IX; C, D, paratype PIN 4270/2540, pupa of male: C, segments V–IX, D, segments VIII–IX.</p> <p> <b>FIGURE 7.</b> <i>Jurochlus lineatus</i> <b>sp. nov.</b>, Shar Teg, J3: A, holotype PIN 4270/2544, pupa of male: segments VIII–IX; B, paratype PIN 4270/348, pupa of male: total view.</p> <p> <b>FIGURE 8.</b> <i>Jurochlus</i> spp. pupae from the Mesozoic of Mongolia: A, <i>J. limbatus</i> <b>sp. nov.</b>, Shar Teg, J3, holotype PIN 4270/ 2584, female: total view (segments VIII–IX); B, C, <i>J. adustus</i> <b>sp. nov.</b>, Khutel Khara, J3/K1, holotype PIN 3965/3209, female: B, total view (pigmentation of tergites not shown), C, segments VIII–IX.</p> <p> <b>Description.</b> Pupa of male. Segment VII slightly widening distally. Each segment with three wide nearly parallel (broadly oval) concavities, a median and two lateral ones, more darkened than surrounding areas. Tergite VII with a well-developed ridge. Lateral margin of segment VII with at least an apical seta. Tergite VIII with posterior margin slightly darkened.</p> <p>Segment IX. Outer margins of anal lobes not sinuate: nearly straight in the middle part and very slightly concave at extreme apex. Outer margin with serration strong, developed distal to base of second seta and reaching apical one-fifth, consisting of 40–50 elongate-triangular teeth; inner margin serration finer, no more than 20 teeth distinct. Segment with strong apical spurs (outlines much impressed into/projecting over the surface).</p> <p> <b>Measurements</b> (holotype, paratype). Combined length of abdominal segments VII–IX, 2800, 2950; entire body length (extrapolated), 9500–10500. Abdominal segments VII–VIII: length, 760, 820–840; width, 1880–1950. Abdominal segment IX: length, 1280–1290; width at base, 1600–1620; distance between tips of anal lobe spurs, 685, 600.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on pages 441-446, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

Jurochlus rigor Kalugina 1985

<i>Jurochlus rigor</i> Kalugina, 1985 <p>(Fig. 1 C)</p> <p> <b>Type material.</b> <i>Holotype</i>: PIN 3053/774 (abdominal segments III–IX of female pupa, Siberia, Uda, Middle or Upper Jurassic).</p> <p> <b>Additions to description.</b> Upon re-examination of the holotype we confirm that no spines, setae, or their attachment places are visible at lateral margins of any segment; anterior and posterior margins are darkened as in <i>J. sibiricus</i>, but details are less clearly visible; genital sac not visible; probable genital plate IX visible near the anterior margin of the segment IX as an impressed transverse stripe with lateral parts slightly curved anteriorly.</p> <p>Anal lobes of the segment IX with inner margins nearly straight, outer margins very slightly sinuate; both darkened, with stout dark apical spurs. Outer margin in distal half with short thick denticles.</p> <p> <b>Measurements.</b> Visible part of abdomen (III–IX) length, 3580; entire body length (extrapolated), 5000–6000. Abdominal segments length/width: V–VII, 500/1250–1380; VIII, 450/1050. Abdominal segment IX: length, 550; width at base, 830; distance between apices of anal lobes, 280.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on pages 438-439, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

Jurochlus limbatus Lukashevich & Przhiboro, 2012, sp. nov.

<i>Jurochlus limbatus</i> sp. nov. <p>(Figs 1 D, 3F–H, 8A)</p> <p> <b>Type material.</b> <i>Holotype:</i> PIN 4270/2584 (abdominal segments VIII–IX of female pupa). SW Mongolia, Gobi- Altai Aymag, Shar Teg (outcrop 443/1), Upper Jurassic.</p> <p> <b>Etymology.</b> From the Latin <i>limbus</i> (fringe), in reference to fine serration of the anal lobe.</p> <p> <b>Description.</b> Pupa of female. Tergite VIII with submedian darkened areas, anterior margin well-sclerotized, posterior margin darkened; five LS setae, broken off, attachment places distinct. Segment IX. Outer margin of anal lobe sinuate, distinctly convex in middle part and strongly concave in apical third. Outer margin finely serrate; serration clearly visible in middle third of margin, consisting of elongate seta-like spines; inner margin without visible serration. No lateral setae visible. Anal lobes apically tapered into sharp darkened spurs; distance between apical spurs 0.35x as long as width at base of segment IX. Genital sac more darkened than outer adjoining area, elongate, apically slightly tapering with truncate posterior margins, with sheaths not rounded apically. Segment IX with well-darkened area anterior and anterolaterally to genital sac (probable genital plate IX).</p> <p> <b>Measurements.</b> Combined length of abdominal segments VIII–IX, 2300; entire body length (extrapolated), 10500–11500. Abdominal segments VIII: length, 900; width, 2100. Abdominal segment IX: length, 1400; width at base, 1750; distance between tips of anal lobe spurs, 600.</p>Published as part of <i>Lukashevich, Elena D. & Przhiboro, Andrey A., 2012, Pupae of Mesozoic Jurochlus Kalugina, 1985 (Diptera: Chironomidae), with description of four new species, pp. 434-452 in Zootaxa 3478</i> on pages 439-441, DOI: <a href="http://zenodo.org/record/282306">10.5281/zenodo.282306</a&gt

A new genus and new species of felt scales (Homoptera: Coccinea: Eriococcidae) from Tierra del Fuego (Chile)

Telmatococcus igniumterrae gen. nov. et sp. nov. is described and illustrated based on the material collected from a sphagnum bog in the extreme southern part of South America (Tierra del Fuego). The new monotypic genus differs from all known genera of Eriococcidae in having short cylindrical truncated setae of the anal apparatus and pouches with numerous quinquelocular pores near the anal apparatus. The habitat of the new taxon is characterized