Calofulcinia oxynota La Greca, 1969

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 : 343-349

Calofulcinia oxynota La Greca

Figures 4–7 , 11A

Calofulcinia oxynota 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 [633].

Type material. The holotype female could not be located by Balderson ( Balderson 1984); it is not in La Greca’s collection, nor is it in the Natural History Museum, London. La Greca’s (1969) detailed description and figures leave no doubt as to the identity of the species, however. He gives the following details for the type specimen:

“ QUEENSLAND: Cairns, Barrow River , 12.IX.1966, 1 ♀ (A. Walford-Huggins leg.) (tipo).”

“Barrow River” is almost certainly misspelt and refers to the Barron River in Cairns.

Specimens examined. QLD: 1♁ Stewart Ck., 4km NNE Mt Spurgeon (Camp 1) 1250–1300m, 16°24’S 145°13’E, 15–20 OCT 1991, Monteith, Janetzki, Cook & Roberts ( QM); 1♁ The Bluff , 11km W of Mossman 800– 1300m, 2 NOV 1983, Monteith, Yeates & Thompson ( QM); 1♁ Mossman Bluff Track 10km W Mossman 1200m, 17 DEC 1988, Monteith & Thompson, pyrethrum/trees & logs ( QM); 1♁ Mossman Bluff Track, 5–10km W. Mossman, Site 8 1180m, 20 DEC 1989 – 15 JAN 1990, Monteith, Thompson & ANZSES, flt. Intercept ( QM); 1♁ Mossman Bluff Summit 10km W Mossman 1300m, 21 DEC 1989, Monteith, Thompson & ANZSES ( QM); 1♁ 2km ESE Mossman Bluff, 9km W Mossman 1000m, 17–18 DEC 1988, Monteith & Thompson ( QM); 1♁ Mt Lewis 500m past swamp, 29 DEC 1989, J. Hasenpusch ( MV); 1♁ Mt Lewis , 23 DEC 1995, J. Hasenpusch ( ANIC); 1♀ Bakers Blue Mt 17 km W Mt Molloy 900m, 11 SEP 1981, G. Monteith & D. Cook ( QM); 1♁ same locality and collectors, 1100m, 12 SEP 1981 ( QM); 1♀ same locality, 800–1000m, 30 DEC 1989 – 9 JAN 1990, ANZES Expedition ( QM); 1♀ Mt Formartine South , 10km N. Kuranda 700m, 23 NOV 1990, Monteith & Thompson, pyrethrum/trees & logs ( QM); 1♁ Kuranda (335m) (Top of the Range) 19 Butler Dr 16.48’S 145.38’E. ( GPS), 16–31 OCT 2008, DCF Rentz, ANIC Database No. 11 000022 ( ANIC); 2♁♁ same locality and collector, 18–30 SEP 2009, ANIC Database No. 11 000023 ( ANIC); 1♀ same locality and collector, 1–15 OCT 2009 ( ANIC); 1♁ same locality and collector, 16–31 JAN 2011, ANIC Database No. 11 000020 ( ANIC); 1♁ same locality and collector, 16–31 MAR 2011, ANIC Database No. 11 000021 ( ANIC); 1♁ same locality and collector, 1–15 FEB 2011, ANIC Database No. 11 000032 ( ANIC); 1♁ same locality and collector, 1–15 AUG 2016 ( ANIC); 1♁ same locality and collector, 16–30 SEP 2016 ( ANIC); 2♁♁ same locality and collector, 1–15 NOV 2016 ( ANIC); 1♁ same locality and collector, 16–31 JAN 2017 ( ANIC); 1♁ same locality and collector, 16–30 APR 2017 ( ANIC); 1♁ same locality and collector, 1–15 JUN 2017 ( ANIC); 1♁ same locality and collector, 16–30 NOV 2017 ( ANIC); 1♁ same locality and collector, 1–15 JAN 2018 ( ANIC); 1♁ same locality and collector, 1–15 FEB 2018 ( ANIC); 2♁♁ same locality and collector, 1–15 MAR 2018 ( ANIC); 1♁ same locality and collector, 1–15 APR 2018 ( ANIC); 1♁ same locality and collector, 1–15 JUL 2018 ( ANIC); 1♁ same locality and collector, 16–31 OCT 2018 ( ANIC); 2♁♁ same locality and collector, 16–30 NOV 2018 ( ANIC); 1♁ same locality and collector, 1–15 JAN 2019 ( ANIC); 1♁ same locality and collector, 16–31 JAN 2019 ( ANIC); 1♁ same locality and collector, 1–15 FEB 2019 ( ANIC); 1♁ same locality and collector, 16–28 FEB 2019 ( ANIC); 1♁ same locality and collector, 1–15 SEP 2019 ( ANIC); 1♁ same locality and collector, 16–31 OCT 2019 ( ANIC); 1♁ same locality and collector, 1–15 MAY 2020 ( ANIC); 1♁ same locality and collector, 16–31 OCT 2020 ( ANIC); 3♁♁ same locality and collector, 16–31 DEC 2020 ( ANIC); 2♁♁ same locality and collector, 16–31 JAN 2021 ( ANIC); 1♁ same locality and collector, 16–28 FEB 2021 ( ANIC); 1♁ same locality and collector, 16–30 NOV 2021 ( ANIC); 1♁ same locality and collector, 1–15 DEC 2021 ( ANIC); 1♁ same locality and collector, 16–31 DEC 2021 ( ANIC); 1♁ same locality and collector, 1–15 FEB 2022 ( ANIC); 2♁♁ same locality and collector, 1–15 MAR 2022 ( ANIC); 1♁ Whitfield Ra., SW. of Cairns 520m, 14 DEC 1970, J.G. Brooks, at light ( ANIC); 1♁ Isley Hills 1050m, 17°03’S 145°42’E, 30 NOV 1993, Cook, Monteith & Janetzki ( QM); 1♀ nymph 1♁ North Bell Peak , Malbon Thompson Ra. 800–1000m, 19–22 NOV 1990, Monteith & Thompson ( QM); 1♀ Nth. Bell Peak, 20 km S Cairns 900–1000m, 15–16 SEP 1981, G. Monteith & D. Cook ( QM); 1♀ with ootheca, Lake Eacham 17.17°S 145.38°E, 8 FEB 1989, G. Milledge, MAN-44 ( MV); 1♁ Lake Barrine , Atherton Tableland, 17.15 S 145.38 E, 11 DEC 1985, J. Balderson, Stop A-8 ( ANIC); 1♀ Massey Range , 4km W of Centre Bellenden Ker 1250m, 17°16’S 145°49’E, 9–11 OCT 1991, Monteith, Janetzki & Cook ( QM); 2♁♁ Mt Baldy Loop Rd , nr Atherton Herberton Range 1097m, 17.16’S 145.25’E. ( GPS), 20 MAR 2010, D.C.F. Rentz & B. Richardson, Stop 3, ANIC Image Database No. 11 000024–000025 ( ANIC); 1♁ Mt Baldy , 17.27 S 145.42 E, 23 MAR 2022, M.G. Connors, C. Henderson & M. Allan, at lights ( ANIC); 1♀ same locality, date, and collectors, on tree trunk with lichen ( ANIC); 1♀ nymph, same locality, date, and collectors, on tree trunk with lichen, raised in captivity, died 11 MAY 2022 ( ANIC); 1♀ same locality, date, and collectors, on tree trunk with lichen, raised in captivity, died 19 NOV 2022 ( ANIC); 1♀ Curtain Fig , 2km SSW. of Yungaburra, Atherton Tableland, 17.17 S 145.34 E, 13 DEC 1985, J. Balderson ( ANIC); 1♁ Wongabel St For. , forest track off Pavey Rd 746m, 17°19.112 S 145°30.315 E (car), 26 NOV 2019, D.C.F. Rentz, Stop 28 ( ANIC); 2♀♀ nymphs, Wongabel , 17.332 S 145.503 E, 30 APR 2022, M.G. Connors, C. Henderson & M. Allan, on tree trunk with moss ( ANIC); 1♀ with ootheca, same locality, date, and collectors, collected as nymph and raised in captivity, matured JUL 2022 ( ANIC); 1♁ 9km W. of Crawfords Lookout (1 road km S. of Palmerston Hwy.), Palmerston Nat. Park, 17.35 S 145.43 E, 11 FEB 1988, D.C.F. Rentz, at light, Stop A-16 ( ANIC); 1♀ Mt. Fisher 7km SW Millaa Millaa ( Kjellberg Rd ) 1000m, 3 MAY 1983, G.B. Monteith & D.K. Yeates ( QM); 1♀ Palmerston Nat. Park, 12km SE. of Millaa Millaa, 17.35 S 145.42 E, 15 DEC 1985, J. Balderson ( ANIC); 1♀ Palmerston Nat. Pk. 350–400m, 2 JAN 1990, G.B. Monteith ( QM); 1♀ Hinchinbrook Is , Upper Gayundah Ck 850m, 10 NOV 1984, G. Monteith, pyrethrum knockdown ( QM); 1♁ Paluma , 1 OCT 1979, D. Frith, light trap in rainforest, Genitalia prep. MG335 J. Balderson ( ANIC); 1♁ same locality and collector, DEC 1980, at light ( ANIC); 1♁ Mt Halifax summit, 21 MAR – 10 MAY 1991, D. Cook, heath, pitfalls & intercepts ( QM); 1♁ Mt Elliott summit 1150m, 26 MAR – 12 MAY 1991, D. Cook, rainforest pitfall & intercepts ( QM) .

Differential diagnosis. C. oxynota can be distinguished from C. australis and C. paraoxypila by the keeled dorsomedian lobes on the third to sixth abdominal tergites of the male abdomen reaching approximately one fifth the length of the following tergite, by the expanded but short, flattened dorsomedian lobes on the third to fifth tergites of the female abdomen, and by the L4A of the male genitalia possessing a dextrally-directed, rounded lobe on the right side. Additionally, C. oxynota can be distinguished from C. australis by the paired tubercles dorsolaterally on the anterior metazone, and it can be distinguished from C. paraoxypila by the slender body form, lack of a postocellar process, and short female tegmina that almost reach the hind margin of the first abdominal tergite.

Description.

Head. Head slightly longer in female. Clypeus with anterior area depressed; anterior margin slightly sinuate in male, more sinuate in female. Lower frons generally six-sided, five-sided in some females. Eyes distinctly projecting dorsally in female. Ocellar tubercle prominent, weakly raised in male, in female low and indistinct; in male ocelli moderately large, ovate, median ocellus distinctly smaller than lateral ocelli, in female all ocelli very small, almost absent. Vertex without obvious postocellar process but broadly raised behind ocelli, weakly concave in dorsal view; juxtaocular bulge weak in male, not strongly projecting beyond vertex, in female stronger, projecting beyond eyes and vertex ( Figure 5D–E ). Antennae reaching beyond midpoint of pronotum in female.

Thorax. Pronotum elongate, somewhat slender, approximately 2.3–2.5 times as long as wide in male, approximately 2.0–2.3 times as long as wide in female, broadest just posterior to supracoxal sulcus. Median keel well-defined on almost entire metazone, generally slightly stronger in female. Prozone ovate, with sides almost parallel, anteriorly with a broad elevated tubercle in male, in female moderately elevated anteriorly; posteriorly with low, undulating diagonal ridges that extend onto lateral prozone. Metazone elongate, approximately 1.7 times length of prozone, anterodorsally with a pair of widely-spaced dorsolaterally-projecting tubercles, these stronger in female, dorsolateral longitudinal carinae rather long, covering posterior half except extreme posterior region, posterior tubercle with a pair of blunt, dorsally-directed spines or rounded knobs in male, in female with a pair of elongate, blunted, dorsally-directed spines. Posterolateral expansion usually acutely-angled and somewhat expanded adjacent to anterior corners of metazone ( Figure 5A–B , 6A–B ). Postcervical sclerite with anterior margin broadly incised; T-shaped sclerite somewhat narrow medially.

Foreleg spination formula: F = 3DS/7–12AvS/4PvS; T = 4–8AvS/4–7PvS.

A single aberrant but apparently undamaged individual from Lake Barrine has only 3 posteroventral forefemoral spines and a strongly reduced middle discoidal spine on the right forefemur.

Legs. Forecoxa very elongate, narrow. Forefemur elongate, narrow, much more so in male, broadest slightly distal to tibial spur groove, concave dorsally; posterior keel very weak, sometimes almost absent; tibial spur groove weakly-defined distally; anteroventral forefemur spines alternating between small and large, with the distal four spines iiiI, IiiI, or iIiI. Foretibia slender.

Wings. Tegmina of male with media occasionally branched; pterostigma slightly raised ( Figure 4B ). Tegmina of female very short, almost reaching hind margin of first abdominal tergite; veins heavily reticulate; pterostigma distinctly raised ( Figure 7A,C–D ). Hind wings of female very short, not exceeding tegmina.

Abdomen. Male abdomen with posterior margin of second to ninth tergite with a short keeled median spine extending posteriorly, in second, seventh, and eighth tergites very small, in other tergites extending up to a fifth of the distance along the next tergite. Female abdomen with second to eighth tergites with dorsomedian and dorsolateral projections, ninth tergite with dorsomedian projection only; dorsolateral projections present as short, blunted, keeled spines, these especially well-developed on third and fourth tergite; dorsomedian projections present as blunted, keeled spines projecting dorsally on second to sixth tergite, projecting posteriorly on seventh to ninth tergite, these distinctly larger, broader, and somewhat laterally compressed on second to fifth tergite, especially so on third and fourth tergite ( Figure 5C ). Cerci somewhat laterally compressed; apical segment small, flattened ( Figure 5F–G ). Supra-anal plate of male with lateral margins broadly convex, exceeding posterior edge of subgenital plate. Supra-anal plate of female similar but larger, acutely produced and slightly blunted posteriorly, exceeding tip of subgenital plate. Subgenital plate of male broadly incised at posterior tip, with styli attachment point within this incision, with a blunted triangular projection on either side of incision; styli short, subconical ( Figure 5G ).

Genitalia. Male genitalia with L4A with a broad, flat, rounded lobe medially on the right side projecting dextrally; pda strongly bifurcate into two flattened, blunted, finely shagrinate triangular lobes, the right lobe directed posterodextrally and moderately bent dorsally, the left lobe rather blunter, directed posteriorly, sometimes very weakly bent dorsally or ventrally; afa with posterior lobe rather elongate, the anterior lobe thicker, broadly rounded, directed anterodistally; paa sometimes very weakly bent to the left; fda moderately broad; pia a very low, concave ridge with one or both corners raised into very small angular projections; pva rather short and robust, projecting ventrally and weakly curving posteriorly, strongly sclerotised, slightly broadening distally and then abruptly narrowing to a blunt point very weakly directed dextrally ( Figure 4C–D ).

Colour. Body colour green, greenish brown, or rarely greenish blue with darker markings, pattern relatively consistent but somewhat variably in intensity and shade.

Head mostly pale green or greenish brown with some dark markings; a pair of narrow, diagonal dark stripes on the lower frons running from the lateral corners to the anterior margin, this often extending slightly onto genae, generally darker in male; a dark posterior margin surrounding the median ocellus, this stronger in female; with very variable dark markings on the vertex, genae, and sometimes lower frons. Antennae annulated along entire length. Eyes green or brown, sometimes very vibrant, with irregular reddish-brown patterning, this fading after death.

Pronotum ground colour relatively pale with variable dark mottling; a very dark spot on the anterolateral margin of the metazone, this sometimes greatly expanded onto the dorsal metazone and lateral pronotal expansion; all dorsal tubercles and a narrow median line pale; lateral pronotal expansion usually with irregularly spaced dark spots; pronotal markings otherwise very variable; green specimens usually weakly marked, browner specimens often strongly patterned with dark spots and blotches. Prosternum mostly pale with longitudinal dark stripes on the postcervical sclerite.

Legs pale with broad mid to dark green or brown bands except on ventral two-thirds of forefemur, often with darker and paler spots in pale regions, anterior surface of forefemur strongly mottled with dark patches; forecoxa and forefemur often with very dark spots along dorsal and posterior keels; first tarsomere usually with basal and median dark band, subsequent tarsomeres indistinctly banded.

Tegmina of male pale greenish, sometimes blueish or almost white; with sparse irregular darker brown or green mottling on plicatum and between branches of anterior cubitus; a broad dark arc in mediodistal quarter of tegmen, running along media and then curving anteriorly to meet anterior radius, in some specimens this marking interrupted medially into two disjunct ovate patches; irregular dark patches on membrane adjacent to distal portions of major veins; sometimes also with irregular dark patches elsewhere; pterostigma pale; most major veins green, posterior cubitus brown, radius and costa with regularly-spaced dark patches, these extending onto membrane; crossveins in pale regions brown, in dark regions green. Tegmina of female pale with some dark mottling basal to pterostigma; an irregular dark spot apically; veins green. Hind wings of male mostly hyaline; costal region and extreme apical portion of remigium pale, opaque; costal crossveins with distinct dark spots where they meet the anterior margin of the wing; membrane adjacent to extreme apical portion of posterior radius, media, and anterior cubitus with irregular dark patches; veins green or pale brown. Hind wings of female almost entirely hyaline; veins pale brown.

Abdomen green and brown, mottled with irregular darker and lighter patches; in female generally brown ventrally, green dorsally, with irregular dark markings dorsolaterally, sometimes with a row of paired pale blue spots on tergites, if present then these especially strong on third to fifth tergites, sometimes also with pale blue patches on dorsolateral expansions; female often with a distinct pale patch on sixth to eighth tergite between dorsomedian and dorsolateral projections; sternites irregularly and faintly mottled; cerci with faint pale annulations ( Figure 4A–B , 7 ).

Nymph. Older nymphs are similar in morphology and colour to adults but lack wings; in females the abdomen is more slender ( Figure 7F ). Very young nymphs are rather long-legged with very small abdominal lobes.

Ootheca. Small, rather robust; externally covered with a thin layer of pale orangish beige foam, internally a darker orangish brown; taller than wide, with lateral sides almost parallel, slightly bulging, weakly converging dorsally, with vertical ridges corresponding to egg chambers; proximal end rather truncate, weakly slanting; distal end almost parallel with proximal end such that the dorsal surface overhangs the base, broadly rounded basally, produced dorsally into an elongate residual process, this approximately one-third the length of the remainder of the ootheca; external foam layer thin but covering almost entire ootheca; emergence area with seven rounded openings in two rows offset from each other, these covered with foam and almost entirely flat ( Figure 6 ). The only known oothecae were deposited in captivity; they are attached to the substrate primarily at the proximoventral end, but one shows signs of weak attachment along the entire ventral surface and the second is weakly attached across most of the ventral surface. In nature, it is probably affixed to bark in a similar manner.

Measurements. Body length, 12.4–17.6 (♀), 14.0–15.8 (♁). Pronotum length, 3.5–5.0 (♀), 2.8–4.9 (♁). Pronotum width, 1.7–2.1 (♀), 1.2–2.2 (♁). Tegmen length, 3.0–3.5 (♀), 14.1–15.3 (♁).

Distribution. C. oxynota has been collected from rainforest and wet sclerophyll forest in northeastern Queensland, primarily at high elevations. It is known from Mount Lewis in the north to Mount Elliot in the south ( Figure 11A , 12A–C ).

Remarks. This is probably the most well-known species of Calofulcinia , although females especially are still very infrequently encountered in comparison to other fulciniine genera. Populations are often very localised, and it is not unusual to collect several males at a single location whilst recording none in other ostensibly suitable areas. Similarly, multiple females or nymphs may inhabit a single tree trunk or rock.

C. oxynota are typically encountered in humid, high-elevation (above 300m) rainforests and adjacent habitat, where they live amongst mosses and lichens. They are especially widespread at localities above 500m elevation, and targeted searches for the species in lowland areas have repeatedly failed to find any specimens.

C. oxynota females have been observed undergoing minor changes in colouration to better match their surroundings. In particular, a captive female collected from Mt Baldy was initially lichen-coloured, but developed a series of teal spots along its abdomen in response to the growth of a similarly-coloured mould in its habitat ( Figure 7C ). When this mould was removed, these spots disappeared. Greater changes in colour and pattern may well be possible, but have not yet been observed.

The hunting behaviour of captive C. oxynota , and to some extent the hunting behaviour of wild C. oxynota , is well-known and the general hunting strategy is detailed under the generic description.Adult males hunt infrequently and generally attempt to catch only small prey, whereas adult females and nymphs of both sexes may take relatively large prey. They appear to notice potential prey well before it is within striking distance, and often wait for it to approach them before pursuing it. Remarkably, we observed an adult female responding to a stimulus more than 2.5 metres away.

Figures

FIGURE 4. Calofulcinia oxynota La Greca.A. Dorsal habitus, subadult female. B. Dorsal habitus, adult male. C. Male genitalic complex, dorsal view. D. Male genitalic complex, ventral view. A–B. Scale = 10 mm. C–D. Scale = 0.5 mm.

FIGURE 5. Calofulcinia oxynota La Greca. 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 6. Calofulcinia oxynota La Greca, ootheca. A. Dorsal view. B. Lateral view. Scale = 1 mm. Photographs by Ken Walker.

FIGURE 7. Calofulcinia oxynota La Greca, live photographs. A. Adult female, Kuranda, Queensland, photograph by Maurice Allan. B. Adult male, Kuranda, Queensland. C. Adult female, Mount Baldy, Queensland, photograph by Maurice Allan; note blue spots. D. Adult female, Kuranda, Queensland, photograph by Maurice Allan. E. Adult male, Lake Eacham, Queensland, photograph by Maurice Allan. F. Female nymph, Mount Baldy, Queensland.

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.