Calofulcinia paraoxypila (Tindale)

Connors, Matthew G., Yeeles, Peter, Lach, Lori & Rentz, David C. F., 2023, Revision of the genus Calofulcinia Giglio-Tos (Mantodea: Nanomantidae: Fulciniinae) in Australia, Zootaxa 5296 (3), pp. 333-361 : 349-357

Calofulcinia paraoxypila (Tindale)

Figures 8–11

Rawarena paraoxypila Tindale, N.B. 1930 , Mantidae in the Australian Museum. Records of the Australian Museum, vol. 17(8), pp. 343–347 [345].

Calofulcinia paraoxypila (Tindale) . La Greca, M. 1969, Su alcuni Mantodei dell’Australia e delle Filippine. Annali del Museo civico di storia naturale Giacomo Doria, vol. 77, pp. 633–639 [636].

Type material. Holotype female. Label 1 front. “K55943 Rawarena paraoxypila Tindale ♀ TYPE”. Label 1 back. “NB Tindale 1. 1929.”. Label 2. “Ulong, East Dorrigo. N.S.W. W. Heron II–IV.1923.”. Label 3. “K55943”. Label 4. “Australian Museum K.55943”. Holotype in the Australian Museum, Sydney .

Allotype male. Label 1. “Ulong, East Dorrigo , N.S.W. W. Heron II–IV·1923 ”. Label 2. “I.18314 Rawarena paraoxypila Tindale ♁ Dorrigo Dist. ALLOTYPE♁”. Label 3. “K49697.” Label 4. “SAMA Database No. 11- 000064”. Allotype in the South Australian Museum , Adelaide .

Paratype male nymph. Label 1 front. “ Rawarena paraoxypila Tindale larva. PARATYPE”. Label 1 back. NB Tindale 1. 1929.”. Label 2. “Ulong, East Dorrigo. N.S.W. W. Heron II–IV.1923 ”. Label 3. “K55944”. Label 4. “Australian Museum K.55944”. Paratype in the Australian Museum, Sydney .

Specimens examined. QLD: 1♁ Lake Barrine , Atherton Tableland, 17.15 S 145.38 E, 11 DEC 1985, J. Balderson ( ANIC) ; 1♁ Curtain Fig ( CSIRO Tower , 35m), 2km SSW. of Yungaburra, Atherton Tableland, 17.17 S 145.34 E, 4 MAR 1988, D.C.F. Rentz, Stop A-2 ( ANIC) ; 1♁ Lake Eacham , Atherton Tableland, 17.17 S 145.39 E, 21 JAN 1988, D.C.F. Rentz, at light, Stop A-3 ( ANIC) ; 1♁ Wongabel (arboretum) (Kennedy Hwy) QLD. 780m, 17°19’39.41’’S 145°30’20.43’’E (GE), 26 APR 2011, D.C.F. Rentz & B. Richardson, Stop 18 ( ANIC) ; 1♁ Broken River , Eungella Nat.Pk., 18–19 APR 1979, G.B. Monteith ( QM) ; 1♁ Broken R., Eungella , 750m, 21°10’S 148°31’E 16–17 NOV 1992, Monteith, Thompson, Cook & Janetzki ( QM) ; 1♀ nymph, Witta 4552, 26.7143 S 152.8151 E, 4 APR 2022, L. Grenfell, iNaturalist observation 110336772 ( ANIC) ; 1♀ Mt Nebo 540m, 27°24’S 152°47’E, end of SEP 2008, G. Karour ( QM) ; 1♁ Davidson’s property hmstd, Tamborine Mtn, 27°57’24’’S 153°11’17’’E, 14 JAN 1993, K.J. Lambkin, at light ( QM) ; 1♁ St. Bernard’s Mt. Tamborine , 28 FEB 1960, C.W. Frazier, At light, ANIC Uni. of New England Coll. Donated 1983 ( ANIC) ; 1♁ Upper Dalrymple Ck , via Goomburra, 21–22 NOV 1987, G.B. Monteith ( QM) ; 1♁ 5km S. of Canungra , 3 DEC 1991, G.B. Monteith ( QM) ; 1♁ “ Glen Witheren ” 5kms S. Canungra, 2–4 JAN 1993, G.B. Monteith ( QM) ; 4♁♁ ‘ Glen Witheren’ , via Canungra 120m, 28°03’S 153°07’E, 19–20 JAN 2000, G. Monteith, at MV light, 9450 ( QM) ; 1♁ Glen Witheren 140m, 28°03.0’S 153°07.2’E, 2–3 NOV 2007, G.B. Monteith, MV light, 15400 ( QM) .

NSW: 1♁ Blue Gum Lookout, via Mallanganee, 7 FEB 1978, G.B. Monteith, Ex UQIC donated 2011 ( QM); 1♁ Moonpah State Forest , via Dorrigo, 11 DEC 1971, G.B. Monteith, Ex UQIC donated 2011 ( QM) .

Differential diagnosis. C. paraoxypila can be distinguished from C. australis and C. oxynota by the small but prominent postocellar process on the vertex, by the longer female tegmina that exceed the hind margin of the second abdominal tergite, by the lack of obvious dorsomedian lobes on the tergites of the male abdomen, by the short, triangular dorsomedian lobes on the third to fifth abdominal tergites of the female abdomen, and by the L4A of the male genitalia possessing a dorsally-directed, rounded lobe on the right side and a posteroventrally-directed lobe on the left side.

Description.

Head. Head slightly longer in female. Clypeus with anterior area weakly depressed; anterior margin slightly sinuate; anterior longitudinal sulcus strong. Lower frons six-sided. Eyes weakly projecting dorsally in female. Vertex almost straight in dorsal view; in male with postocellar process present as a distinct, strongly raised, rounded tubercle; in female entire anterior vertex strongly raised, with a blunt triangular projection directed anterodorsally; juxtaocular bulge moderate in male, weakly projecting beyond vertex, in female stronger, projecting well beyond vertex ( Figure 9D–E ). Antennae reaching almost to posterior end of pronotum in female.

Thorax. Pronotum relatively short, broad, approximately 1.7–2.2 times as long as wide in both sexes, broadest at supracoxal sulcus. Median keel weakly-defined to well-defined on almost entire metazone, usually slightly weaker in female, generally weaker in northern specimens. Prozone short, almost circular, anteriorly with a very low tubercle in male, in female moderately elevated anteriorly; posterolaterally with low diagonal ridges, these stronger in male. Metazone relatively short, approximately 1.9 times length of prozone, elevated posteriorly, anterolateral sulcus rather shallow, anterodorsally with a pair of low, widely-spaced dorsolaterally-projecting tubercles, these stronger in female, dorsolateral longitudinal carinae rather short, covering posterior third except extreme posterior region, posterior tubercle with a pair of blunt, dorsolaterally-directed tubercles. Posterolateral expansion somewhat expanded and angulate adjacent to anterior corners of metazone ( Figure 8A–B , 9A–B ). Postcervical sclerite with anterior margin almost straight in male, weakly concave in female; T-shaped sclerite narrow medially.

Foreleg spination formula: F = 3DS/10–11AvS/4PvS; T = 7–8AvS/5–6PvS.

Legs. Forecoxa elongate, slightly narrowing distally. Forefemur somewhat broad, robust, broadest at tibial spur groove, with scattered tubercles, slightly concave dorsally, more so in female; dorsal keel much weaker in female; posterior keel very weak in male, sometimes almost absent, in female virtually absent; tibial spur groove moderately-defined distally; anteroventral forefemur spines alternating between small and large, with the distal three spines iiI or IiI. Foretibia robust.

Wings. Tegmina of male with veins somewhat reticulate, with dense crossveins only in extreme basal region of remigium ( Figure 8B ). Tegmina of female short, weakly saddle-shaped such that the distal end is slightly raised, just exceeding posterior margin of second abdominal tergite ( Figure 8A ). Hind wings of female short, slightly exceeding tegmina.

Abdomen. Male abdomen with posterior margin of tergites weakly convex, with a short median keel on posterior section. Female with second to ninth tergites with dorsomedian and dorsolateral projections; dorsolateral projections present as short, blunted, keeled spines, these especially well-developed on third tergite; dorsomedian projections present as blunted, keeled spines projecting dorsally on third to fifth tergite, projecting posteriorly on sixth to ninth tergite, these distinctly larger, broader, and somewhat laterally compressed on third to fifth tergite; third and fourth tergites with a pair of distinct rounded tubercles adjacent to dorsomedian projections ( Figure 9C ). Cerci very weakly laterally compressed; apical segment elongate, cylindrical ( Figure 9F–G ). Supra-anal plate of male with lateral margins strongly convex, not reaching posterior edge of subgenital plate. Supra-anal plate of female ovate, with median keel almost absent, broadly rounded posteriorly, not reaching tip of subgenital plate. Subgenital plate of male broadly incised to rounded at posterior tip, with a short triangular lobe dorsally to each stylus, these lobes sometimes very small, rarely absent; styli short, subconical, setose, often asymmetrical such that one of the styli is significantly shorter and more conical than the other ( Figure 9G ). Subgenital plate of female with ventroterminal lobes somewhat elongate ( Figure 9F ).

Genitalia. Male genitalia with L4A with a rounded, shagrinate, dorsally-directed lobe medially on the right side, with a more elongate, rounded, posteroventrally-directed lobe posteromedially on the left side; pda very small, very finely shagrinate, rounded to subtriangular, not strongly sclerotised, directed almost dextrally; afa with posterior lobe elongate, slightly tapering to a blunt point, directed posterodextrally but curved slightly such that the tip is directed dextrally, the anterior lobe much shorter and broader, very low, sometimes virtually absent; paa very weakly bent to the left; L4B with a very small, flattened, sinistrally-directed lobe posteromedially on the left side; fda rather broad, sometimes slightly constricted basally; pia a very low, rather flat ridge, sometimes with one or both corners raised into very small angular projections; pva rather short and robust, projecting ventrally and weakly curving posteriorly, slightly broadening distally and then abruptly narrowing to a blunt point very weakly directed dextrally ( Figure 8C–D ).

Colour. Body colour very variable, pattern relatively consistent but variable in intensity. Most individuals vaguely to diffusely lichen-coloured, i.e. very pale green or greenish blue with black or dark brown markings, some individuals with pale areas almost white, other individuals brown or greenish brown with more diffuse dark markings and strong mottling, some brown individuals almost entirely without dark areas ( Figure 8A–B , 10 ). The following description of colour is based primarily on lichen-coloured individuals ( Figure 8A–B , 10B,D,F ).

Head mostly pale, in male with scattered dark spots and blotches, usually with a transverse row of dark spots on lower frons, ocellar tubercle often strongly mottled, with a transverse line extending across dorsal surface of head, interrupted by ocelli, this line sometimes not extending onto genae, often with curved dark lines on vertex adjacent to parietal sulcus. Female head almost entirely pale, ocellar tubercle dark, with a solid dark transverse line extending from lateral ocellus almost to margin of eye; posterior region of vertex strongly mottled in browner individuals. Antennae strongly annulated along entire length, with dark annulations gradually widening distally. Eyes similar in colour to head, spotted, usually with a dark lateral stripe extending across the entire length of the eye, these markings fading after death.

Pronotum ground colour similar to head, with a dark spot anterolaterally on the metazone, this sometimes extending onto dorsal surface of metazone, onto lateral pronotal expansion, and/or onto lateral region of prozone; in some lichen-coloured specimens also with a sinuate dark border on the posterolateral metazone; in mottled specimens usually with a narrow, diffuse greenish border along the entire pronotum; lateral pronotal expansion with scattered dark spots adjacent to prozone and posterior of metazone. Prosternum mostly pale with irregular and variable dark markings.

Legs mostly pale with darker markings. Forecoxa with large dark blotches, in some females these conjoined into three dark bands; forefemur anteriorly with four large dark blotches, narrowly or broadly conjoined, the distalmost two or three of these extending dorsally and onto the posterior surface, these marks paler and more diffuse on posterior surface of mottled individuals; foretibia banded on anterior surface, banded or without pattern on posterior surface. Mid and hind legs with regular dark bands, sometimes absent basally on femora, paler regions often with mottling. Tarsomeres of all legs distally dark, first tarsomere also with a medial dark band.

Tegmina of male pale green or greenish blue with dark brown markings, often with strong brown spotting or mottling; base of tegmen medially dark, pterostigma almost white, a small dark wedge distal to pterostigma; membrane adjacent to medial section of media dark, forming an elongate dark marking, this sometimes broken into two smaller dark marks; membrane adjacent to posterior radius with a dark patch where it meets the anterior radius, from this a dark marking extending posteromedially such that the entire dark area is subtriangular, with concave edges; membrane adjacent to extreme distal regions of sectors irregularly darkened; with irregular dark markings between first anterior cubitus and plicatulum; radius and costa with regularly-spaced dark spots along almost their entire length, these denser on the costa, often also extending onto adjacent membrane. Veins mostly pale cream, dark in shaded regions of tegmen, posterior cubitus brown. Tegmina of female with pterostigma almost white, with a dark ovate diagonal spot of similar size at the distal and apical end of the pterostigma; in lichen-coloured females, almost the entire rest of the tegmen concolorous with pterostigma; in mottled females more irregularly marked with spots and dark patches; veins usually concolorous with membrane, in mottled females costal region with veins darker, greenish. Hind wings of male mostly hyaline; costal region and extreme apical portion of remigium pale, opaque; costal crossveins distally with distinct dark spots where they meet the anterior margin of the wing; membrane adjacent to extreme apical portion of posterior radius, media, anterior cubitus, and sometimes anterior branch of second anterior analis with irregular dark patches; veins pale brown. Hind wings of female mostly hyaline, extreme apical portion similar in colour to tegmen, veins mostly pale brown.

Abdomen similar in colour to thorax, very variable in pattern. In lichen-coloured females, mostly pale with irregular, dark, diagonal lines on second to fourth tergites, these sometimes extending onto dorsomedial projections; usually also with irregular dark patch on seventh tergite; in males and mottled females, abdomen very irregularly patterned, generally greenish dorsally and brown ventrally, sometimes with large pale spots laterally on third to seventh tergites. Cerci unicolorous or with faint dark annulations ( Figure 8A–B , 10 ).

Nymph. Older nymphs are similar in morphology and colour to adults but lack wings; in females the abdomen is more slender ( Figure 10C–D ). Very young nymphs are rather long-legged and frequently have bolder patterning than adults and larger nymphs.

Ootheca. Unknown but possibly similar to that of C. oxynota .

Measurements. Body length, 14.4–15.9 (♀), 13.9–17.5 (♁). Pronotum length, 3.7–4.4 (♀), 3.3–4.2 (♁). Pronotum width, 1.8–2.3 (♀), 1.7–2.5 (♁). Tegmen length, 4.0–4.2 (♀), 13.8–14.3 (♁). 9.87755

Distribution. C. paraoxypila is known from a very broad latitudinal range in eastern Queensland and northeastern New South Wales from a variety of forested habitats. It is widespread between Eumundi in the north and Taree in the south, but specimens have also been collected from much further north, at Eungella and on the Atherton Tablelands ( Figure 11 , 12A–B, D, F–H ).

Remarks. This robust, stocky species is the most widespread of the Australian Calofulcinia , and in the southern portion of its range it is probably the most frequently-encountered. It has been collected from a variety of habitats ranging from undisturbed rainforest to suburbia, and has been collected from moss- or lichen-covered tree trunks, rocks, and even bare rock and paperbark tree trunks.

Perhaps because of this broad habitat and microhabitat preference, it is also the most variable species in both colour and pattern. Although most males are relatively similar to each other, females may show a variety of different patterns ( Figure 8A 10A, E ). This enables them to effectively camouflage themselves against a variety of different substrates, although their ability to change their colouration as C. oxynota does is untested. This colour variation is far greater in the southern portion of its range than in the north; only lichen-coloured individuals have been recorded from the northern portion of its range.

Despite its relative abundance in southeast Queensland and northern New South Wales, C. paraoxypila is only very rarely collected in north Queensland, where it is known from just four male specimens and two photographic records. These were collected at lights in dense, relatively high-elevation (above 500m) rainforest, suggesting that the species may display a markedly different life history strategy in these tropical regions. It is possible that this population usually lives high in the canopy, as is suspected to be the case for many other uncommon insects known from the area ( Hasenpusch & Brock 2006; Rentz et al. 2012).

This species is rather dissimilar to the other Australian members of the genus and to all of the Papuan species, being larger and very much more robust than their delicate congeners. Indeed, Tindale (1930) initially described it in the monotypic genus Rawarena . He noted that it was “allied to Stenomantis and to Ima ” (all Australian species of Stenomantis have now been transferred to Ciulfina ) but did not compare it to any of the numerous Papuan genera of Fulciniini which were at that point considered valid, but most of which are now considered synonyms. La Greca (1966) expressed doubt at the validity of Rawarena , and later synonymised it with Calofulcinia , finding that the differences between C. paraoxypila and C. oxynota were not warranting of generic status ( La Greca 1969).

While this synonymy is accepted here, a more thorough review of the entire Fulciniini is required before any definitive statements on generic boundaries can be made. At present it seems unlikely that Rawarena should be considered a distinct genus, but future studies may find that it is warranting of subgeneric status.

Figures

FIGURE 8. Calofulcinia paraoxypila (Tindale).A. Dorsal habitus, adult female. B. Dorsal habitus, adult male. C. Male genitalic complex, dorsal view. D. Male genitalic complex, ventral view. A–B. Scale = 10mm. C–D. Scale = 1 mm.

FIGURE 9. Calofulcinia paraoxypila (Tindale). A. Lateral pronotum, adult female. B. Lateral pronotum, adult male. C. Lateral abdomen, adult female. D. Head, adult female. E. Head, adult male. Subgenital plate, adult female. F. Subgenital plate, adult male. A–B, D–F. Scale = 1 mm. C. Scale = 5 mm.

FIGURE 10. Calofulciniaparaoxypila (Tindale), livephotographs.A.Adultfemale,mottled morph,Carneys Creek,Queensland, photograph by Greg Neill. B. Adult male, diffuse lichen morph, Mapleton, Queensland, photograph by Dianne Clarke. C. Female nymph, patternless morph, Coffs Harbour, New South Wales, photographs by Nick Lambert. D. Nymph, lichen morph, Witta, Queensland, photograph by Larney Grenfell. E. Adult female, green morph, Repton, New South Wales, photograph by Jeannie Bartram. F. Adult male, diffuse lichen morph, Mapleton, Queensland, photograph by Dianne Clarke.

FIGURE 11. Distribution of Calofulcinia spp. in Australia. A. Distribution in northeastern Queensland. B. Distribution in southeastern Queensland and northern New South Wales. Green = C. australis, Blue = C. oxynota, Purple = C. paraoxypila. Stars = type locality, Circles = specimens, Triangles = photographic records. Base map courtesy of Google Maps ©2023 and TerraMetrics ©2023.

FIGURE 12. Typical habitat of Calofulcinia spp. A. Rainforest at Lake Eacham, Queensland, habitat of C. oxynota and C. paraoxypila, photograph by Matthew Borella. B. Rainforest edge at Wongabel, Queensland, habitat of C. oxynota and C. paraoxypila. C. Transitional rainforest-wet sclerophyll forest at Mount Baldy, Queensland, habitat of C. oxynota. D. Rainforest at Tamborine Mountain, Queensland, habitat of C. australis and C. paraoxypila, photograph by Caitlin Henderson. E. Rainforest at Cunninghams Gap, Queensland, habitat of C. australis, photograph by John-Michiel Koens. F. Rainforest at Guanaba, Queensland, habitat of C. australis and C. paraoxypila, photograph by Nicholas John Fisher. G. Wet sclerophyll forest at Coffs Harbour, New South Wales, habitat of C. paraoxypila, photograph by Nick Lambert. H. Rural garden/restored rainforest edge at Witta, Queensland, habitat of C. paraoxypila, photograph by Larney Grenfell.