Description of a new species of Aoplonema (hemiptera: miridae: orthotylinae) with a biogeographic analysis and reassessment of the phylogeny of the genus.
|Publication:||Name: Entomologica Americana Publisher: New York Entomological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Science and technology Copyright: COPYRIGHT 2009 New York Entomological Society ISSN: 1947-5136|
|Issue:||Date: Jan, 2009 Source Volume: 115 Source Issue: 1|
|Topic:||Event Code: 310 Science & research|
|Geographic:||Geographic Scope: Mexico Geographic Code: 1MEX Mexico|
Abstract--Aoplonema lopidium sp. nov. is described from the Mexican
states of Guerrero and Oaxaca, representing the first records of this
genus from mainland Mexico. A summary of the hypothesized phylogenetic
species relationship is (A. princeps (A. lopidium (A. rubrum + A.
nigrum))). A biogeographical analysis proposes that the Sierra Madre del
Sur is the sister area to the rest of the areas occupied by the other
species of Aoplonema.
Key words: Heteroptera, Aoplonema lopidum, biogeography, Mexico.
The genus Aoplonema Knight belongs to a monophyletic group of Orthotylini genera denominated the Hadronema group (Forero, 2008). Seven genera are placed in the Hadronema genusgroup, all of which, among other characters, share the presence of a supragenital bridge (Konstantinov, 2003:11), and the particular structure of the right paramere, which is hatchet-shaped with a long body and a dorsomedial flattened apical process (Forero, 2008: 15). Aoplonema has three described species, A. nigrum Forero, A. princeps (Uhler), and A. rubrum Forero. These species uniquely possess a single sclerotized vesical spicule, which is deeply cleft in the middle (Forero, 2008: 153). Known species of Aoplonema are distributed across North America, and have the southern limit of their distribution in northern Baja California, Mexico.
Recently, when Forero's (2008) paper was in press, the junior author realized that among material from the Texas A&M entomological collection, there was an additional, undescribed species of Aoplonema, which was not noted before because of the strikingly different dorsal coloration. In this paper, we describe this new species of Aoplonema from southern Mexico, provide a new key to the species of Aoplonema, assess the phylogenetic position of this colorful new species within Aoplonema, and provide a biogeographical analysis.
MATERIALS AND METHODS
Scanning electron micrographs of gold-coated and uncoated preparations were taken with a digital Philips XL30 ESEM. Dorsal habitus photographs were taken on a Microptics-USA photographic system equipped with an Infinity Photo-Optical K-2, three-lens system and Nikon D1X digital camera. All specimens have attached a unique specimen identifier (USI) number, which has the format AMNH_PBI followed by the identification number (e.g., AMNH_PBI 00234987). Each USI uniquely identifies a particular specimen from the Plant Bug Planetary Biodiversity Inventory project (http://research.amnh.org/pbi/). Additional information can be retrieved with the USI number via the Discover Life website (http://www.discoverlife. org/).
Morphological terms for male and female genitalia follow Davis (1955), Kelton (1959), Scudder (1959), Konstantinov (2003), and Forero (2008). Other morphological terms follow Schuh and Slater (1995), except in the term "collum" that refers to the posterior part of the cephalic capsule posterior to the transverse basal carina (see e.g., Knight, 1941). All measurements are in millimetres unless otherwise stated. In the "material examined" section, interpolated data--in particular georeference--are between square brackets.
For the phylogenetic analysis all four species of Aoplonema, including A. lopidium, were treated as terminals of the ingroup. In a previous phylogenetic analysis (Forero, 2008: 152), Aoplonema was regarded the sister group to the remaining members of the Hadronema group. We selected two members from this clade: Hadronema militare Uhler, 1872, and Scutomiris setosus Forero, 2008, the type species of their respective genera, in order to provide outgroups closely related to Aoplonema. In addition, we included Orthotylus marginalis Reuter, 1883, the type species of the nominal tribe, as an outgroup not related to the Hadronema group, to root the tree. A morphological matrix (Table 1), coded from the vestiture, head, thorax, and male and female genitalia (Table 2), was constructed using WinClada (Nixon, 2002). This matrix was based mostly on the character matrix of the Hadronema group (Forero, 2008: 156) from which uninformative characters pertinent to the terminals included in this matrix of Aoplonema were deleted. In addition, other characters dealing with the particular configuration of the male genitalia of Aoplonema were added to include the variation observed in A. lopidium.
The optimality criterion used in the phylogenetic analysis was parsimony. Tree searches were carried out in NONA (Goloboff, 1999) spawned from WinClada (Nixon, 2002). A heuristic search was carried out holding a maximum of 2001 trees in memory, with 100 replications and 20 trees to hold per replication, in random addition sequence. For each replication a Subtree Pruning and Regrafting and a Tree Bisection and Reconnection swapping were performed (Swofford et al., 1996). Bootstrap and Jackknife values were calculated in WinClada.
A biogeographical analysis was done using the phylogeny obtained. Brooks Parsimony Analysis (BPA) (Wiley, 1988) was used to obtain a fundamental area cladogram. Despite their importance in biogeographic analysis (e.g., Harold and Mooi, 1994; Szumik et al., 2002), areas of endemism for Western North America are not well defined. Some analyses dealing with North America define broad, general areas for which the taxa are coded. We follow Schuh (2006) for the delimitation of some of these areas in Western North America. We thus recognize as areas of endemism Northern California, Southern California, Baja California (the northern part), and the remaining areas of distribution in Western North America as a single area of endemism. None of the species of Aoplonema is restricted to any one of these defined areas, but are found in more than one of these areas. Aoplonema lopidium is found exclusively in the Sierra Madre del Sur, Mexico. This area has been previously recognized as an area of endemism for other insects (Marshall and Liebherr, 2000).
KEY TO THE SPECIES OF AOPLONEMA
The following key is for males only. Female specimens are best identified when associated with males of the same collection event.
1 Eye large, protruding laterally and dorsally (i.e., eye width/interocular distance ratio [greater than or equal to] 0.6) 2
--Eye smaller, not so strongly projecting (i.e., eye; eye width/interocular distance ratio < 0.55) princeps
2 Hemelytron very long, cuneus nearly three times as long as wide; head dark; scutellum red; right portion of the vesica reaching about the middle of the left portion 3
--Hemelytron short (Fig. IA), cuneus less than 2.6 times as long as wide; head white with paired dark markings (Figs. 1A, B); scutellum black with apex orange; right portion of the
vesica reaching the apex of the left portion (Fig. 3) lopidium, new species
3 Hind femur black or dark red with irregular black spots; fight portion of vesica not directed medially ......................... nigrum
--Hind femur red, without black spots; right portion of vesica directed medially ...... rubrum
Aoplonema lopidium, new species
DIAGNOSIS: Recognized by the contrasting orange and black color pattern, including the pale anterior margin of the pronotum, the orange medial stripe on the scutellum, the entirely orange cuneus, the cream colored head with bold black markings, and the mostly black leg, including the basal half of the hind femur (Figs. I A, B); male eye large, reaching the dorsal margin of the head and the short anteocular region (Figs. 2A); both sexes with approximately equal length hemelytra (Figs. 1A, B); right portion vesica with length of the as long as the left portion (Fig. 3), left portion with the diameter of the dorsal rami much narrower than in the ventral rami (Fig. 3); lateral margin of posterior wall with dorsal lobe of the interramal sclerite (= "K" structure of Slater, 1950) gently curved (Fig. 4); and the accessory sclerite on the lateral portion of the dorsal labiate plate laterad to sclerotized ring (Fig. 4).
Because the coloration of A. lopidium is so unusual for the genus, this species was initially not recognized as belonging to the Hadronema group until the male genitalia were examined, revealing a supragenital bridge and particular structure of the parameres, as described below. Thus, the dorsal coloration of both sexes will serve to distinguish A. lopidium from all congeners.
[FIGURE 1 OMITTED]
DESCRIPTION: Male: Delicate, moderately large, total length 3.684.30. COLORATION (Fig. 1A): Head: Pale cream; black on clypeus, most of adjoining maxillary plate, frons ventrally and ringed bilateral parallel stripes, bilateral portions of the vertex near each eye, vertex medially, and collum; labrum dark brown; labium dark brown to black, antenna black. Thorax: Collar, pronorum, mesoscutum, and scutellum black, except pale orange on anterior margin of pronotum, lateral area to callus, and apex and short medial stripe on scutellum; prosternum cream; propleuron pale orange, blending to cream ventrally; mesosternum black; mesepisternum pale to dusky orange; mesepimeron cream; metepisternum cream with dark brown to black dorsally; mesothoracic spiracle and adjacent evaporative area and metepisternal scent gland evaporative area black, peritreme partially pale. Hemelytron: Black except for costal margin, basal half of radius and adjacent corium, and entire cuneus orange; membrane and veins black. Legs: Black except for cream coxa and trochanter, base of coxa black. Abdomen: Dusky orange, with black sublateral band, lateral margin dusky yellow. Genitalia: Genital capsule and parameres black; proctiger dusky cream. STRUCTURE: Head: Strongly declivent; anteocular region short, about a third of head length; frons weakly convex, nearly flat; gula shorter than buccula; eye big, surpassing dorsal margin of head in lateral view, interocular distance small (Fig. 1A). Thorax: Calli nearly flat; mesothoracic spiracle and evaporative area as in Figure 2B. Hemelytron: Each nearly parallel, not greatly exceeding apex of abdomen; cuneus nearly flat, longer than wide (Fig. 1A). Legs: Pretarsus as in Figure 2C. Genitalia: Genital capsule small (Fig. 3); parameres as in Figure 3; left portion of vesica slightly compressed dorsoventrally at apex, denticulate, diameter of ventral ramus wide, heavily denticulate apically, nearly reaching middle of left portion, diameter of dorsal ramus narrow, apex slightly denticulate (Fig. 3); fight portion long, narrow, slightly sinuate, reaching to apex of left portion, slightly denticulate at apex (Fig. 3).
[FIGURE 2 OMITTED]
Female: Similar to male but slightly shorter, ovoid, total length 3.90-4.28. COLORATION: Similar to male (Fig. 1B). Abdomen: Pale yellowish brown with scattered blotches of brown laterally; sternite 9, subgenital plate, and ovipositor dark tan. STRUCTURE: Genitalia: Interramal sclerites and dorsal lobes as in Fig. 4, dorsomedial region of posterior wall finely denticulate; sclerotized rings broadly ovoid, posterior margin medially produced, apex truncate, medial margin smoothly curved, accessory sclerite small, attenuate medially, not reaching interior of sclerotized ring in dorsal view (Fig. 4); anterior wall membranous, and inner margins of first gonapophyses symmetrical (Fig. 4).
MEASUREMENTS: Male (n=3 unless noted otherwise; mean followed in parentheses by range): Length, 4.08 (3.68-4.30); clypeus to cuneus 2.67 (2.45-2.85); width, 1.29 (1.14-1.45). Head length, 0.42 (0.404).44); width through eyes, 0.88 (0.830.90); vertex width, 0.29 (0.284).31). Length of antennal segment I, 0.29 (0.284).30); II, 1.23 (1.14-1.30); III, 0.96 (0.85-1.05); IV (n=2), 0.34 (0.33-0.34). Labium length, 1.05 (0.99-1.10). Pronotal length, 0.65 (0.604).69); width, 1.09 (1.00-1.14). Scutellum length, 0.45 (0.41-0.48); width 0.54 (0.50-0.56). Cuneal length, 0.64 (0.570.69); width, 0.33 (0.30-0.36).
Female (n=9; unless noted otherwise; mean followed in parentheses by range): Length, 4.1 l (3.90-4.28); clypeus to cuneus 2.79 (2.60-2.90); width, 1.32 (1.25-1.43). Head length, 0.42 (0.4-10.45); width through eyes, 0.86 (0.834).90); vertex width, 0.44 (0.414).45). Length of antennal segment I, 0.27 (0.254).28); II, 0.89 (0.854).95); HI (n=7), 0.70 (0.63-0.78); IV (n=7), 0.35 (0.310.38). Labium length, 1.11 (1.06-1.19). Pronotal length, 0.70 (0.66-0.76); width, 1.24 (1.20-1.31). Scutellum length, 0.47 (0.44-0.51); width 0.62 (0.60-0.63). Cuneal length, 0.58 (0.544).61); width, 0.34 (0.30-0.36).
[FIGURE 3 OMITTED]
DISTRIBUTION: Aoplonema lopidium is known from three localities in the Sierra Madre del Sur, with the distribution thus far localized in the northern and central regions of Guerrero and southeastern Oaxaca, Mexico (Fig. 5).
HOSTS: No host plant data is noted on the specimen labels.
ETYMOLOGY: Named to indicate the superficial similarity of the color pattern to small species of the genus Lopidea Uhler, 1872.
DISCUSSION: The new species is the only member of the genus thus far known from southern Mexico. Other species of Aoplonema are distributed in North America from the 100th Meridian to the West coast, and from Canada to Baja California, Mexico (Forero, 2008). In both sexes, the contrasting orange and black dorsal color pattern, black legs, and pale cream-colored head with bold black markings are unique to the genus. J. C. Schaffner (pers. comm.), the collector of all specimens of A. lopidium, also collected a least three undescribed species of Lopidea from the Cacahuamilpa area that have a color pattern similar to A. lopidium.
HOLOTYPE MALE: MEXICO: Guerrero: 2.1 mi N of Cacahuamilpa, [18.67339[degrees]N 99.55333[degrees]W], 19 Jul 1984, Carroll, Schaffner, Friedlander, l[male] (AMNH_PBI 00245399) (UNAM).
PARATYPES: MEXICO: Guerrero: 2.1 mi N of Cacahuamilpa, [18.67339[degrees]N 99.55333[degrees]W], 19 Jul 1984, Carroll, Schaffner, Friedlander, 1[female] (AMNH_PBI 00245400) (TAMU). 2.1 mi NW of Cacahuamilpa, [18.67484[degrees]N 99.5602[degrees]W], 11 Aug 1978, Plitt and Schaffner, l[male] (AMNH_PBI 00245392) (AMNH), 1[female] (AMNH_PBI 00245393) (TAMU); 23 Jul 1981, Bogar, Schaffner and Friedlander, 1[male] (AMNH_PBI 00245394), 4[female] (AMNH_PBI 00245395-AMNH_PBI 00245398) (TAMU); 10 Aug 1980, Schaffner, Weaver, Friedlander, 1[female] (AMNH_PBI 00245402) (AMNH). 6mi E of Xochipala, [17.7936[degrees]N 99.54263[degrees]W, 1,067 m], 13 Jul 1985, Jones and Schaffner, 1[female] (AMNH_PBI 00245403) (TAMU). Oaxaca: 8 mi SE of El Camaron, [16.50105[degrees] N 95.19749[degrees]W, 1,356 m], 19 Jul 1987, Kovarik and Schaffner, 1[female] (AMNH_PBI 00245401) (TAMU).
[FIGURE 4 OMITTED]
As a result of the analysis we obtained one tree with a length of 34 steps, consistency index of 79, and retention index of 66 (Fig. 6). Only unambiguous characters are mapped onto the tree obtained (Fig. 6).
Aoplonema lopidium is considered a member of the Hadronema group because of the presence of a supragenital bridge (character 9), which is weakly sclerotized (character 10), and the structure of the right paramere (characters 12-14) (Fig. 3) (Forero, 2008). Because the tree obtained in this analysis (Fig. 6) shows only unambiguous optimizations for the characters, and because there is only one non-Hadronema group species present as an outgroup (i.e., O. marginalis), the characters supporting the inclusion of A. Iopidium in the Hadronema group are not shown in Figure 6.
[FIGURE 5 OMITTED]
Aoplonema lopidium is further considered to belong into Aoplonema because the vestiture is composed of two types of setae: simple decumbent and sericeous (characters 0-1) (Fig. 2D); the phallotheca is cylindrical, with the opening mostly dorsal, and with a basal enlargement on the left side (characters 15 17) (Fig. 3); and the vesica has a single spicule (character 18), which is deeply cleft, and is formed of two portions, with the left portion apically with two cephalad-directed rami (Fig. 3). In our analysis (Fig. 6), A. lopidium is the sister species of A. rubrum + A. nigrum based on the short anteocular region of the head (character 2) (Fig. 2A), and the large eyes surpassing the dorsal margin of the head in lateral view (character 4) (Fig. 2A). Males of A. lopidium do not share with A. rubrum and A. nigrum the elongate hemelytron and cuneus (character 7) (Fig. 1A).
The Sierra Madre del Sur, which is the area occupied by A. lopidium, is resolved as the sister area of the remaining areas occupied by the other species of Aoplanema (Fig. 7). Of the four species of Aaplanema, only A. lapidium occupies such a restricted distributional area. Other species of Aaplanema occupy different geographical ranges in North America, mostly restricted to Western North America (Forero, 2008: 49). Other orthotyline taxa in the Orthatylus genus-group (see Schuh, 1974) have similarly restricted distributions to the Sierra Madre del Sur; for example, the genus Ficinus Distant, many of the species of Jornandes Distant (Schaffner and Schwartz, 2008), several new species of Sealponotatus Kelton and Lopidella Knight, and a new species of a new genus near Jornandes (M. D. Schwartz, pers. obs.). This is in agreement with previous studies for treating this area as an area of endemism (e.g., Liebherr, 1994).
The Sierra Madre del Sur has been proposed as the sister area of the Sierra Transvolcanica in Mexico, and both areas are nested within a larger clade of central and southern Mexican areas. This group of areas is considered sister to the remaining areas to the north, including several areas in the southwestern United States (Marshall and Liebherr, 2000).
[FIGURE 7 OMITTED]
Because A. lopidium is not the most basal taxon in the phylogeny, it can be argued that an extinction event, or a yet not discovered species of Aoplonema, may be responsible for the incongruent pattern observed. Further collecting efforts in Southern Mexico will certainly reveal additional species that can be used to evaluate this biogeographical hypothesis.
This project was supported by, and represents a contribution to, the National Science Foundation Planetary Biodiversity Inventory grant DEB-0316495 to Randall T. Schuh (AMNH) and Gerasimos Cassis, University of New South Wales, Sydney, Australia.
Thomas J. Henry and two anonymous reviewers greatly improved the manuscript with comments and suggestions. We especially thank Joseph C. Shaffner, who collected all the specimens of A. lopidium, and made them available to us. Acronyms, names of institutions, and curators or individuals responsible of the collections mentioned in the text are listed below:
AMNH American Museum of Natural History, New York, Randall T. Schuh
UNAM Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico D.F., Harry Brailovsky
TAMU Texas A&M University, College Station, Joseph C. Schaffner and Edward C. Riley
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Received and accepted December 31, 2008.
D. FORERO (1,2) AND M. D. SCHWARTZ (1)
(1) Division of Invertebrate Zoology, American Museum of Natural History, New York, New York 10024, USA. e-mail: email@example.com
(2) Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853, USA. e-mail: firstname.lastname@example.org
Table 1. Character matrix. Character Taxon 00 01 02 03 04 05 06 07 08 09 Ortkotylus marginalis 0 0 0 0 0 0 1 0 1 0 Hadronema militare 1 0 0 0 0 0 1 0 1 1 Scutomiris setosus 1 1 0 1 0 1 0 0 0 1 Aoplonema lopidium 0 1 1 0 1 0 1 0 0 1 Aoplonema princeps 0 1 0 0 0 0 1 0 0 1 Aoplonema rubrum 0 1 1 0 1 0 1 1 0 1 Aoplonema nigrum 0 1 1 0 1 0 1 1 0 1 Character Taxon 10 11 12 13 14 15 16 17 18 19 Ortkotylus marginalis - 0 0 0 0 0 0 0 0 - Hadronema militare 0 0 1 1 1 0 1 1 0 - Scutomiris setosus 0 1 1 1 1 1 1 1 0 - Aoplonema lopidium 1 1 1 1 1 1 0 0 1 1 Aoplonema princeps 1 1 1 1 1 1 0 0 1 0 Aoplonema rubrum 1 1 1 1 1 1 0 0 1 0 Aoplonema nigrum 1 1 1 1 1 1 0 0 1 0 Character Taxon 20 21 22 23 24 25 26 Ortkotylus marginalis - - 0 0 1 0 1 Hadronema militare - - 1 0 0 1 0 Scutomiris setosus - - 1 0 0 0 1 Aoplonema lopidium 1 0 0 0 1 0 0 Aoplonema princeps 0 0 0 1 1 0 0 Aoplonema rubrum 0 0 0 1 1 0 1 Aoplonema nigrum 0 1 0 1 1 0 1 Table 2. Character descriptions. Vestiture: 0. Simple setae on hemelytron: decumbent = 0; erect = 1. 1. Sericeous setae on hemelytron: not present = 0; present = 1. Head: 2. Anteocular region of head in male: about one-third the length of head = 0; less than one-third the length of head 3. Vertex, posterior margin: with transverse carina = 0; nearly flat, not elevated = 1. 4. Eye in lateral view: reaching dorsal margin = 0; surpassing dorsal margin = 1. Thorax: 5. Disc of scutellum in lateral view: flat = 0; swollen = 1. 6. Metepisternum vestiture: sparse macrotrichia = 0; dense macrotrichia = 1. 7. Cuneus, relative length in male: less than 2.6 times as long as wide = 0; about three times as long as wide = 1. Male Genitalia: 8. Apex of genital capsule: subtriangular = 0; subquadrangular = 1. 9. Supragenital bridge: absent = 0; present = 1. 10. Supragenital bridge, degree of sclerotization: strong = 0; weak = 1. 11. Sclerotized part of ductus seminis: not elongate = 0; elongate = 1. 12. Right paramere dorsal apophysis: not medially directed = 0; medially directed = 1. 13. Right paramere at apical half: not fiat = 0; flattened = 1. 14. Shape of right paramere in medial view: gently curved = 0; hatchet-shaped = 1. 15. Phallotheca: without protuberances = 0; with a lateral protuberance on basal left side = 1. 16. Opening of phallotheca, shape: mostly dorsal, with parallel margins = 0; reclined, oval = 1. 17. Proximoventral surface of phallotheca: well sclerotized = 0; weakly sclerotized = 1. 18. Number of spicules: 2 = 0; 1 = 1. 19. Single spicule, right portion, length relative to left portion: about half the length = 0; about the same length = 1. 20. Single spicule, right portion, shape of apex: curved upward = 0; slightly curved downward = 1. 21. Single spicule, rami, relative length in respect to total length of spicule: about half = 0; about one third = 1. Female Genitalia: 22. Dorsal lobe of interramal sclerite, medial margin: sub-apically enlarged = 0; nearly straight, slightly sinuate = 1. 23. Dorsal lobe of interramal sclerite, lateral margin: curved = 0; sub-apically enlarged = 1. 24. Dorsal lobe of interramal sclerite, apex: narrowing distally but not acute = 0; rounded = 1. 25. Anterior wall: membranous = 0; with a sclerotized central plate = 1. 26. Accessory sclerite of sclerotized ring, relative position: lateral to sclerotized ring = 0; recurved over sclerotized ring = 1.
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