Cremnorrhinina Reuter

Randall T. Schuh & Michael D. Schwartz, 2016, Nineteen new genera and 82 new species of Cremnorrhinina from Australia, including analyses of host relationships and distributions (Insecta: Hemiptera: Miridae: Phylininae: Cremnorrhinini), Bulletin of the American Museum of Natural History 401, pp. 1-279 : 6-31

Cremnorrhinina Reuter

Cremnorrhinaria Reuter, 1883: 567 (descr.).

Cremnorrhinini Wagner, 1974: 367 (descr.). Cremnorrhini [sic] Schuh and Menard, 2013: 21 (status); Menard et al., 2014: 407 (com position and relationships).

Cremnorrhina [sic] Schuh and Menard, 2013: 21 (as subtribe); Menard et al., 2014: 407 (composition and relationships).

DIAGNOSIS: Recognized by the elongate pulvilli (fig. 31E, F), in the Australian fauna unique in always having these structures free from the claw except at the base and extending to the apex of the claw. Usually with moderate sexual dimorphism, female shorter and more strongly ovoid than male, weakly to strongly brachypterous. Coloration ranging from almost entirely pale (pls. 8, 28), sometimes nearly white (pl. 2, 16), to almost completely black (pl. 16, 24); tibial spines usually with pale bases (e.g., pl. 12), much less frequently with dark bases (pl. 16). Head sometimes strongly prognathous and elongate (pl. 26), although usually with face short to very short (e.g., pls. 22, 28). Head below eyes never more than one-half height of head to as little as one-fifth (figs. 8B, 45B). Antenna usually with segments 2, 3, and 4 slender, 3 and 4 of more or less equal diameter (e.g., pls. 26, 28), segment 2 occasionally swollen and terete (pl. 24); antennal sexual dimorphism weak, segment 2 in male sometimes more robust than in female. Pronotum usually flattened, trapezoidal, lateral margins ranging from weakly concave to weakly convex, posterior lobe sometimes moderately swollen and elevated; calli ranging from inconspicuous to distinctly demarcated; mesoscutum narrowly to broadly exposed; scutellum in form of equilateral triangle, flattened or very weakly elevated. Male genitalia typical for the Phylinae in structure of phallotheca, left and right parameres; endosoma, formed of two distinct straps, ranging from very slender and long to short and compact, J-shaped or sigmoid, sometimes with one or two lateral spines arising near secondary gonopore, apex with one, two, or three spines of various shapes and lengths and sometimes with a membranous baglike structure, frequently ornamented with microtrichia or denticles. Structure of female genitalia typical for phylines with subgenital plate of sternite 6 variable, usually concave medially, sometimes with posteriorly

directed medial projection; vestibular sclerites, formed by bases of first gonapophyses and ventromedial extension of ventral labiate plate, usually a compact subsymmetrical structure, rarely large and asymmetrical; dorsal labiate plate with variable size and shape of sclerotized rings; posterior wall simple, usually composed of two lateral sclerites and one medial sclerite, rarely lateral sclerites joined dorsomedially; intersegmental membrane between sternites 8 and 9 sometimes with discrete structure.

DISCUSSION: Schuh and Menard (2013) and Menard et al. (2014) rendered the tribal and subtribal names of this group as Cremnorrhini/Cremnorrhina, an obvious lapsus. We here employ the orthographically correct renderings as used by Reuter (1883), Wagner (1974), and others.

Definitive recognition of cremnorrhinine taxa in Australia is often possible only through inspection of the pretarsus, because the habitus can be remarkably similar to that found in other tribal-level groupings. Nonetheless, as we demonstrate through the use of scanning electron microscopy, the structure of the pretarsus is essentially monotonous across the entire group and unique within the fauna (figs. 8F, 31E, F, 45F, 57D), and therefore an unequivocal character. A very limited number of Australian phylines, all of them currently undescribed, have enlarged pulvilli that are adnate to the entire ventral surface of the claw, but in those cases the claws are shorter than in the Cremnorrhinina. Although the analyses of Menard et al. (2014) and Schuh and Menard (2013) placed some taxa from the Holarctic with this type of claw structure in the Cremnorrhinina (e.g., Pronotocrepis Knight , Dacota Uhler ), many other taxa with such enlarged pulvilli, including the few from Australia, do not appear closely related to the Cremnorrhinina, and in the latter case the structure of the male genitalia also supports their exclusion. Many such examples from the Northern Hemisphere belong to the Nasocorini (e.g., Atractotomus Fieber , Rhinacloa Reuter , among many others), with a few, such as Eminoculus Schuh (Hallodapini) from South Africa, belong to other tribal-level groupings.

We offer additional commentary on relationships and comparisons with cremnorrhinine taxa from other zoogeographical areas in the General Discussion section at the end of this paper.

The following key to genera will serve as an additional aid to recognition of the genera we propose in this paper and assist in the organization of material for the identification of species. The key will be most effective when used to identify male specimens, because many of the characters used in the formation of our generic concepts, as well as for species recognition, are derived from the male genitalia. Positive identification of many genera will require dissection of the male genitalia as is the case for most species. For a large number of genera, identification of females will be possible only through association with males.

KEY TO GENERA OF AUSTRALIAN

CREMNORRHININA

1. Dorsum with contrasting dark spots at bases of setae (pls. 1, 2, 16, 18)................. 2

– Dorsum frequently with some dark markings, e.g., at base of cuneus or on membrane, but never at bases of setae (e.g., pls. 6, 12, 24).. 6

2. Vestiture a mixture of dark, reclining common setae and flattened, weakly lepidote setae (pl. 18)...................... Lepidophylus

– Vestiture with only simple setae.......... 3

3. Body form relatively broad; spots on dorsum forming blotches with ill-defined margins (pl. 16); endosoma with two, somewhat twisted, apical spines of subequal length; secondary gonopore with large denticles on outer surface (figs. 46, 47, pl. 17)..... Halophylus (in part)

– Body form moderately to strongly elongate; spots on dorsum small, with well-defined margins; endosomal structures not as above.... 4

4. Endosoma elongate, slender, and J-shaped, without easily distinguished straps and no apical spines; secondary gonopore relatively small, compact, subapical in placement (pl. 18)...................... Maculiphylus

– Endosoma sigmoid or J-shaped, with two distinct straps and one or two apical spines; secondary gonopore more elongate or unsclerotized and not visible................... 5

5. Straps of endosoma confluent and fused proximal to secondary gonopore, and forming a single distal strap and an elongate, erect, apical spine (fig. 2, pl. 1); right paramere very large, left paramere boxlike, apical portion of posterior process forming a right angle (fig. 2)....................... Adunatiphylus

– Straps of endosoma distinct over entire length, forming two apical spines; secondary gonopore either normally developed and sclerotized or unsclerotized (figs. 4, 5, pl. 2).. Asterophylus

6. Head weakly to very strongly porrect, often extending beyond anterior margin of eye by distance equal to or greater than length of eye (pl. 26); endosoma with a single hooked apical spine (figs. 76–80, pl. 27)........... Pulvillophylus

– Head not conspicuously porrect, although frons sometimes swollen and clypeus visible from above; apical endosomal spines never hooklike........................... 7

7. Eyes often strongly bulging (pls. 6, 20, 24, 30)................................. 8

– Eyes not so strongly bulging........... 11

8. Small, body form elongate, tubular; head and vertex narrow, eyes bulging; frons often tumid; antennal segments 1 and/or 2 either entirely dark or with some dark areas and not swollen; corium with a dark spot apically (or entirely dark) (pls. 6, 30); endosoma with variable lateral spine near secondary gonopore, usually with microtrichiate surface (pls. 21, 25), and phallotheca sometimes with fields of denticles (figs. 20–22, 24)..................... 9

– Size variable; body not so distinctly tubular; head and vertex wider; frons usually not tumid; antennal segments 1 and/or 2 pale and sometimes conspicuously swollen; corium without dark spot apically (pls. 20, 24); endosoma sometimes with a lateral spine near secondary gonopore, usually without microtrichiate surface (pls. 21, 25), but phallotheca never with fields of denticles (figs. 61–63, 71–75)......... 10

9. Secondary gonopore situated subapically in line with approximately straight shaft of endosoma; endosoma with slender, usually proximally oriented, lateral spine, either subtending or at level of secondary gonopore; phallotheca with one or two fields of denticles on posteroventral surface (figs. 20–25, pl. 7)....... Dicyphylus

– Secondary gonopore situated at midpoint of endosoma, removed laterad of shaft, with flanking crest, and shaft of endosoma strongly bent to the left; endosoma with apically directed narrow spine originating at level of secondary gonopore (figs. 92, 93, pl. 30)............ Telophylus

10. Coloration variable, from almost entirely black to almost entirely pale; antennal segment 2 in male often weakly to strongly swollen, even terete; at least some portion of antenna dark (pl. 24); endosoma with two more or less parallel apical spines, never with a spine at level of secondary gonopore (figs. 71–75, pl. 25)........................ Proteophylus

– Coloration, including antennae, nearly uniformly lime green (pl. 20); endosoma apically bifid, with a single, sometimes bifid lateral spine at level of secondary gonopore (figs. 61–63, pl. 21)....................... Myrtophylus

11. Specimens ranging from mostly brown to mostly pale, but especially distal half of hind femur always with dark spots (pl. 16)................... Halophylus (in part)

– Coloration variable, but femora never with dark spots........................ 12

12. Uniform green or yellow coloration, including appendages, never with areas of infuscation or other dark markings (pl. 19); body robust, especially head and pronotum; dorsum with dark, recumbent simple setae; endosoma sigmoid with two simple apical spines about length of secondary gonopore (figs. 58, 59, pl. 19)...................... Myoporophylus

– Usually mostly green or yellow, but often with some dark markings or infuscate areas on hemelytron; body elongate or with length less than 3.00; dorsal vestiture variable; endosomal structure variable, sometimes similar to form seen in Myoporophylus ............... 13

13. Elongate, relatively large (body length greater than 4.00), with infuscate markings at apex of corium and apex of membrane cells; dorsum covered with dark, recumbent, simple setae (pl. 3); endosoma sigmoid, with two long, weakly curving, more or less parallel apical spines, and a third much shorter spine (pl. 3); posterior process of left paramere elevated (figs. 6, 7).................... Austroplagiognathus

– Coloration often similar to above, but body size usually somewhat smaller (often less than 4.00); vestiture of dorsum variable; structure of male genitalia variable, but never with long parallel spines as in Austroplagiognathus ......... 14

14. Size often relatively small, never elongate (pl. 10) but genitalia large; endosoma short, broad, heavily sclerotized; phallotheca often with a dorsal crest (figs. 32–36, pl. 11)........ Grandivesica

– Size variable, sometimes elongate but endosoma not as short and broad as in Grandivesica .......................... 15 15. Endosoma sigmoid, apically with an inflated membranous bag of variable shape, covered with numerous erect spicules, and with a single lateral spine arising at level of secondary gonopore (figs. 82–91, pl. 29)........ Spinivesica

– Endosoma of variable shape, without an apical, spicule-covered, membranous bag in conjunction with a single lateral spine at level of secondary gonopore...................... 16

16. Body of endosoma twisted, sometimes ropelike, with 2 (or 3) apical spines; membranous covering of endosoma usually evident and with some spicules on apical portion (figs. 37–44, pls. 13, 15)................. Gyrophallus

– Body of endosoma not so strongly twisted as in Gyrophallus and with differing apical spines................................ 17

17. Endosoma with a single robust spine at least as long as or longer than length of secondary gonopore (figs. 54–56, pl. 18); right paramere coming to a distinct point apically (figs. 54– 56)................... Monospiniphallus

– Right paramere with a vaguely to distinctly bifid apex; apex of endosoma with 1 or 2 spines; posterior process of left paramere usually distinctly elevated; if apex of endosoma with a single elongate spine then right paramere broad and distinctly bifid apically............ 18

18. Right paramere with a broad, distinctly bifid apex (figs. 13–18)............ Bifidostylus

– Apex of right paramere not so broad, vaguely bifid............................ 19

19. Endosoma with an elongate apical spine, usually subtended by one or two shorter spines, or sometimes with a single shorter, curving spine (figs. 26–30, pl. 9)........ Eremotylus

– Apex of endosoma with one or two short, slender apical spines (figs. 65–69, pl. 23)...................... Omnivoriphylus

Tables

TABLE 1 Measurements of Australian Cremnorrhinina

			Length						Width			Ratio
			Cun-								AS2/	IOD/ HW/	PW/
Species		Body	Clyp Head	Prono	Scut	Cun	Head	Prono	Scut IntOcDi	AntSeg2	HW	HW PW	BL
Adunatiphylus
A. kalbarri
♂ (N = 5) Mean 3.09 2.00 0.20 0.33 0.30 0.63 0.63 0.80 0.38 0.30 0.88 1.38 0.47 0.80 0.26	Mean	3.09	2.00 0.20	0.33	0.30	0.63	0.63	0.80	0.38 0.30	0.88	1.38	0.47 0.80	0.26
	SD	0.18	0.06 0.02	0.01	0.02	0.04	0.02	0.04	0.02 0.01	0.03	0.05	0.01 0.02	0.01
	Range	0.44	0.15 0.06	0.04	0.04	0.09	0.06	0.10	0.05 0.02	0.07	0.13	0.02 0.05	0.02
	Min	2.79	1.91 0.17	0.32	0.27	0.58	0.60	0.75	0.36 0.29	0.85	1.31	0.46 0.78	0.25
♀ (N = 5)	Max Mean	3.23 2.51	2.06 0.23 1.77 0.23	0.36 0.33	0.31 0.27	0.67 0.42	0.66 0.65	0.85 0.78	0.41 0.31 0.35 0.35	0.92 0.73	1.45 1.12	0.48 0.83 0.54 0.84	0.27 0.31
	SD	0.11	0.06 0.02	0.01	0.01	0.03	0.01	0.01	0.01 0.01	0.03	0.03	0.01 0.01	0.02
	Range	0.30	0.16 0.04	0.02	0.02	0.05	0.02	0.04	0.04 0.03	0.07	0.08	0.04 0.02	0.04
	Min	2.37	1.69 0.21	0.32	0.26	0.39	0.64	0.76	0.33 0.33	0.70	1.09	0.51 0.82	0.29
Asterophylus	Max	2.67	1.85 0.25	0.34	0.28	0.44	0.66	0.80	0.37 0.36	0.77	1.17	0.55 0.84	0.33
A. chrysocephali
♂ (N = 7)	Mean	3.53	2.31 0.23	0.39	0.34	0.67	0.68	0.89	0.45 0.29	1.03	1.50	0.43 0.77	0.25
	SD	0.29	0.13 0.03	0.03	0.03	0.08	0.03	0.06	0.04 0.02	0.08	0.07	0.04 0.01	0.01
	Range	0.83	0.37 0.07	0.09	0.07	0.26	0.10	0.18	0.10 0.05	0.23	0.22	0.12 0.05	0.03
	Min	2.96	2.07 0.20	0.35	0.30	0.50	0.63	0.80	0.39 0.27	0.87	1.38	0.37 0.74	0.24
♀ (N = 7)	Max Mean	3.79 3.17	2.44 0.27 2.17 0.24	0.44 0.39	0.37 0.32	0.76 0.53	0.73 0.68	0.98 0.91	0.49 0.32 0.45 0.35	1.10 0.93	1.60 1.36	0.49 0.79 0.52 0.75	0.27 0.29
	SD	0.16	0.13 0.03	0.03	0.01	0.04	0.03	0.03	0.02 0.01	0.06	0.05	0.02 0.01	0.01
	Range	0.48	0.43 0.09	0.08	0.04	0.12	0.08	0.09	0.06 0.03	0.18	0.13	0.05 0.03	0.02
	Min	2.84	1.91 0.20	0.35	0.30	0.46	0.63	0.85	0.42 0.34	0.82	1.30	0.49 0.73	0.28
A. rutidosis	Max	3.32	2.34 0.29	0.43	0.34	0.58	0.71	0.94	0.48 0.37	1.00	1.43	0.54 0.76	0.30
♂ (N = 7)	Mean	3.12	2.04 0.23	0.34	0.30	0.59	0.66	0.83	0.40 0.30	0.92	1.38	0.46 0.79	0.27
	SD	0.53	0.27 0.02	0.02	0.03	0.13	0.04	0.06	0.04 0.01	0.14	0.14	0.04 0.02	0.03
	Range	1.32	0.67 0.04	0.05	0.08	0.32	0.10	0.14	0.10 0.03	0.37	0.36	0.10 0.05	0.07
	Min	2.50	1.71 0.21	0.31	0.25	0.44	0.61	0.76	0.35 0.29	0.71	1.16	0.42 0.77	0.23
♀ (N = 8)	Max Mean	3.82 2.63	2.38 0.25 1.92 0.25	0.36 0.34	0.33 0.27	0.76 0.41	0.71 0.68	0.90 0.83	0.45 0.32 0.40 0.37	1.08 0.87	1.52 1.26	0.52 0.82 0.55 0.82	0.30 0.32
	SD	0.38	0.20 0.03	0.03	0.02	0.13	0.03	0.05	0.04 0.01	0.05	0.06	0.03 0.04	0.03
	Range	0.97	0.52 0.08	0.08	0.06	0.35	0.09	0.15	0.10 0.03	0.14	0.20	0.09 0.11	0.08
	Min	2.22	1.68 0.21	0.29	0.25	0.26	0.62	0.76	0.36 0.36	0.77	1.12	0.51 0.76	0.28
	Max	3.19	2.20 0.29	0.37	0.31	0.61	0.71	0.91	0.46 0.39	0.91	1.32	0.60 0.87	0.35